www.nature.

com/scientificreports

OPEN Inventory reveals wide biodiversity

of edible insects in the Eastern
Democratic Republic of Congo
Jackson Ishara  1,2*, Rodrigue Ayagirwe3, Katcho Karume4, Gustave N. Mushagalusa4,
David Bugeme5, Saliou Niassy6, Patchimaporn Udomkun  7 & John Kinyuru2

In response to growing food demand, edible insects are perceived as an opportunity to alleviate
food insecurity. With its wide edible insects’ biodiversity, the Democratic Republic of Congo is one
of Africa’s most critical entomophagous. This study aimed at giving a first insight on inventory
showing diversity, perception, consumption, availability, host plants, harvesting techniques and
processing techniques of edible insects in South-Kivu, DRC. It recorded twenty-three edible insects
belonging to nine families and five orders, some of which are consumed in the larval, adult, egg and
pupa stages. Rhyncophorus phoenicis, Alphitobius diaperinus, Macrotermes subhyalinus and Acheta
domesticus were the most preferred edible insects in Fizi Territory, Ruspolia differens and Apis mellifera
larvae in Kabare Territory, Imbrasia oyemensis, Imbrasia epimethea, Rhynchophorus ferrugineus and
Rhyncophorus phoenicis in Mwenga Territory, Ruspolia differens, Macrotermes subhyalinus, Gryllotalpa
africana, Nsike, Nomadacris septemfasciata and A. mellifera larvae in Walungu Territory. Ruspolia
differens, I. oyemensis, A. mellifera larvae, G. africana and Nsike, were preferred for their taste. Acheta
domesticus, A. diaperinus and A. mellifera larvae were abundant throughout the year, while others
were only available for 9 months or less per year. Numerous plants have been recorded as their hosts,
including plants used for food and income. Harvesting strategies and period, processing methods
and preservation techniques depend on insect species, local knowledge and practices. These findings
suggest similar and thorough studies on entomophagy across the country while encouraging the
rearing of edible insects to address their existing high demand and environmental concerns.

The world’s population is expected to reach 9.8 billion by 2050 and 11.2 billion by ­21001, raising concerns about
food production and the ever-growing demand for p ­ rotein2. Edible insects are among the most important biore-
sources being promoted to address global food and nutritional ­security3–5. Worldwide edible insects are regularly
consumed by 2 billion p ­ eople6,7 for their nutritional value and t­ aste8,9. Several studies have demonstrated the
superior nutritional value of edible insects compared to conventional f­ oods10–13.
The most consumed groups of edible insects include beetles (Coleoptera, 31%), caterpillars (Lepidoptera,
18%), and bees, wasps, and ants (Hymenoptera, 14%), followed by grasshoppers, locusts and crickets (Orthop-
tera, 13%), cicadas, leafhoppers, planthoppers, scale insects and true bugs (Hemiptera, 10%), termites (Isoptera,
3%), dragonflies (Odonata, 3%), flies (Diptera, 2%) and 5% other o ­ rders13. The availability of some edible insects
depends on the geographical distribution of their host plants and s­ easonality14 and correlates with their har-
vesting ­period15. In turn, successful edible insect harvest depends on insect habits and ecological f­ actors16, as
they are harvested at different growth stages, including the larval (bees, beetles, butterflies, and ants) and adult
(beetles, ants, grasshoppers) stages. For some species, harvesting is easy at night or early in the morning when
inactive and cannot ­fly17.
In Africa, harvesting techniques of edible insects are a mixture of observations, hand-picking, tracings,
sign interpretations, and trapping s­ trategies18, varying from one insect to another. The most commonly used

1
Department of Food Science and Technology, Université Evangélique en Afrique, P.O. Box  3323, Bukavu,
Democratic Republic of Congo. 2Department of Food Science and Technology, Jomo Kenyatta University of
Agriculture and Technology, P.O. Box 62000‑00200, Nairobi, Kenya. 3Department of Animal Sciences, Université
Evangélique en Afrique, P.O. Box: 3323, Bukavu, Democratic Republic of Congo. 4Department of Crop Sciences,
Université Evangélique en Afrique, P.O. Box: 3323, Bukavu, Democratic Republic of Congo. 5Crop Production and
Protection Unit, Université Catholique de Bukavu, Bukavu, Democratic Republic of Congo. 6International Centre
of Insect Physiology and Ecology (Icipe), P.O. Box 30772‑00100, Nairobi, Kenya. 7International Institute of Tropical
Agriculture (IITA), Bujumbura, Burundi. *email: [email protected]

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 1

Vol.:(0123456789)
 www.nature.com/scientificreports/

Figure 1.  Map showing South-Kivu Province and the study Territories (ArcMap 10.4. https://​deskt​op.​arcgis.​
com/​en/​arcmap/​10.4/).

harvest techniques are hand-picking, mainly for crickets, Rhinoceros beetle, African palm weevil, and caterpil-
lars, while light trapping is the most used technique to harvest termites, green grasshoppers, house cricket, and
mole ­cricket19,20.
Studies on edible insects have been conducted in Africa, America, Asia, Europe and Australia to assess key
variables in entomophagy perception and ­practices5,13,16,21–27. Like many African countries, food security has
been a significant challenge in the Democratic Republic of Congo (DRC), exacerbated by climate c­ hange28,
forcing disadvantaged communities to desperately depend on forest products, including edible insects, as they
are among the most abundant forest r­ esources29,30. They play an incredible role throughout the year, especially
during the rainy season, depending on the geographical location of different c­ ultures31.
Despite the vast diversity of edible insects in the DRC, there are few studies on edible insects’ inventory except
the study conducted on ecological diversity of edible insects and their potential contribution to household food
security in Haut-Katanga P ­ rovince31 and in the area surrounding LuiKotale, Salonga National P ­ ark32. However,
to the best of our knowledge, there is little-to-no research on the biodiversity, perception, consumption, avail-
ability, host plants, harvesting techniques, and processing techniques of edible insects in South Kivu. Studies
from other regions cannot be extrapolated to South-Kivu consumers due to ecological, cultural and dietary habit
differences, meaning there is a need for more detailed study in this regard.

Material and methods
Ethics statement.  All experimental protocols, as well as methods, were approved and carried out as per
relevant guidelines and regulations from the Interdisciplinary Centre for Ethical Research (CIRE) established
by the Evangelical University in Africa, Bukavu, D.R. Congo, with reference (UEA/SGAC/KM 132/2016). All
subjects signed an informed consent form describing the aim of the study after translation into local languages.

Study area.  The survey and direct observation were carried out in four selected Territories (Fizi, Kabare,
Mwenga and Walungu) of South-Kivu Province, D.R. Congo (Fig. 1), where data on diversity, host plants, sea-
sonal availability, harvesting techniques and traditional processing, consumption, preference of edible insects
were collected. These Territories were purposively selected for their familiarity with entomophagy as local com-
munities traditionally practice it. Territories are in different agroecological conditions with different cultures
influencing edible insects’ availability and consumption.

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 2

Vol:.(1234567890)
www.nature.com/scientificreports/

Territory
Characteristics Fizi Walungu Kabare Mwenga
Latitude (South) 3° 30 to 4° 51′ 32 2° 38′ 2° 30′ 3° to 4°
Longitude (East) 27° 45 to 29° 14′ 10 28° 40′ 28° 30′ 28° 25′ 29″
Area ­(km2) 15,789 1800 1960 11,172
Altitude (m) 750 to 1700 1000 to 2000 1420 to 3200 670 to 1800
Humid wet and dry
Climate type Humid wet tropical Humid wet tropical Equatorial
tropical
Ferralsols, Cambisols and
Dominant soil unity Acrisols and Cambisols Ferralsols and Nitisols Acrisols and Cambisols
Nitisols
Mean T °C 23.54 °C 17–20 °C 22.6 °C 21–37 °C
Mean annual P (mm) 1704 900 to 1500 1572 1650
Estimated population
1,093,926 1,509,175 868,616 843,636
(2019)
Density of population
69.3 838.4 443.6 75.5
(hab ­km−2)
AEZ* Low and high altitude Medium to high altitude Medium to high altitude Low and high altitude

Table 1.  Agro-ecological conditions of the study area (retrieved from CAID). P (mm) Precipitation (rainfall),
AEZ Agro-Ecological Zone (High, Medium, Low).

Agro‑ecological conditions of the study area.  The agro-ecological conditions of the study area are
presented in Table 1. The Fizi Territory is located between 3° 30 and 4° 51 32 latitude (South), and 27° 45 and 29°
14 10 longitude (East). Its elevation is subdivided into four zones, including the coastline (~ 750 m), the low land
valley (~ 1000 m), a highland (~ 1300 m), and the very highland (locally called Haut Plateau with 1700 m). The
climate in Fizi is highly affected by the elevation. The rainfalls are unevenly distributed according to the month
and the climatic subdivision. The North, dominated by the coastline and low inland valley, is characterised by
humid tropical climate (of Aw3 type according to Köppen–Geiger classification). The greatest rainfall amounts
are recorded in March and November, while the smallest amounts are the smallest amounts in February and
September. The south part has a dry humid tropical climate. Available climate data mentioned an average annual
rainfall of ~ 1704 mm, the mean temperature ~ 23.54 °C (with the highest observed in April with ~ 25.6 °C and
the lowest ~ 21.3 °C in September). The Territory is dominated by forest, comprising two forest reserves and a
nature reserve. Acrisols and Cambisols are the dominant soil unities according to the WRB classification.
The Kabare Territory is located between 2° 30′ of South latitude and 28° 30′ of East longitude. Its altitude
varies from ~ 1420 to 3200 m, and the Territory occupies an area of ~ 1690 ­km2 with an estimated population
of ~ 868,616, which makes it among the most populated in the South-Kivu province. The Territory is located in the
medium to high altitude AEZ. Available meteorological data mentioned an annual rainfall average of ~ 1572 mm,
and a temperature of ~ 22.6 °C. Most of Kabare is savanna with natural vegetation consisting of wild grasses.
The Mwenga Territory is located in the middle of the province and is the only Territory surrounded by the
other without any country or province borders. It is located between 28° 25′ 29″ East longitude and 30° 02,16′
05″ South latitude. Its altitude varies between 1500 and 1800 m in the northeast. In the centre and the South,
it is more or less 670 m. In the East, it is more or less 200 m and in the West more or less 670 m. It has a humid
tropical climate with two seasons: the dry season from June to September and the rainy season from September
to May. The temperature varies between 21 and 37 °C in most of the Territory and is low in the Itombwe area
because of the high altitude, which goes up to over 2000 m. Rainfall reaches 2000 mm to 3000 mm per year. The
vegetation is mainly dense forest and savanna. The forest is home to the Itombwe Nature Reserve (RNI). Relief
is dominated by the Itombwe mount uplands and the alluvial valley of the Elila watershed. Soils dominated with
clayey (Humic Cambisols) and sandy soil (Acrisols) types.
The Walungu Territory is located between 2° 38′ of South latitude and 28° 40′ of East longitude. Its altitude
varies between 1000 m and 2000 m with a cold tropical climate of low altitude. There are two seasons, the dry
season (June to August) and the rainy season from September to March. Available station data presented an
annual average of ~ 17–20 °C and 900 and 1500 mm for temperature and rainfall, respectively. The vegetation
mainly consists of grassland, a few forest reserves of Mugaba and Mushwere and woodlands scattered throughout
the Territory.

Sampling and selection of respondents.  A total of 520 respondents, about 130 respondents in each
Territory, were interviewed, with priority given to people familiar with entomophagy based on the main objec-
tives set. Therefore, the respondents included adults, women and men over 18  years old and from all social
classes. A structured oral interview was used individually to ensure better information and minimise external
influences on the respondent’s side.

Sources of data collected.  Primary data were obtained from the field survey using three techniques: ques-
tionnaire administration, direct observations, and insect collection.

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 3

Vol.:(0123456789)
 www.nature.com/scientificreports/

Questionnaire administration.  Structured questionnaires were used to obtain information on edible insects
in all selected Territories of South-Kivu. The questionnaire was divided into seven sections. In the first section,
information about the respondents was collected. Section two contained open-ended questions related to com-
monly consumed edible insects, focusing on local names and stages of consumption. The third section contained
questions about consumer preferences and preference factors. The fourth section included questions related to
seasonal availability. In the fifth section, questions about host plants and signs of presence were asked, followed
by personal observations. The sixth section dealt with harvesting techniques and timing. The last section dealt
with processing methods and preservation techniques. The enumerators translated the questions into the local
dialect to enhance the understanding of respondents. Pictures and real samples of various edible insects identi-
fied from the literature were also used to help respondents identify the insects being mentioned. Enumerators
probed further to clarify some responses to enhance the depth of information solicited.

Direct observations.  Direct observations of relevant information related to insects and their habitats in the
different territories were recorded in the field. Pictures were taken to verify and support the responses obtained
from the interviewees. In addition, the researcher had the opportunity to observe how some edible insects were
prepared and consumed.

Collection and taxonomic identification of insect samples.  Samples of edible insects were collected as part of
the survey and were preserved in 70% alcohol before being taken to the laboratory at Lwiro Research Center for
identification. A mixture of primary data and taxonomic characters was used to identify and classify the various
species of edible insects in the different Territories. The taxonomic characters were derived mainly from archival
sources and published literature.

Data analysis.  Data were analysed using R 4.0.0. and Microsoft Excel 16.56. The completed questionnaires
were cleaned and information verified. Based on the nature of the research questions, descriptive and explora-
tory approaches were used to delineate and describe the existence and use of edible insects in various Territories.

Results and discussion
Commonly consumed edible insects in selected Territories of South‑Kivu.  A total of twenty-
three edible insects including Macrotermes subhyalinus, Acheta domesticus, Rhyncophorus phoenicis, Alphitobius
diaperinus, Ruspolia differens, Gryllotalpa africana, Apis mellifera larvae, Nomadacris septemfasciata, Locusta
migratoria, Rhynchophorus ferrugineus, Imbrasia oyemensis, Imbrasia epimethea, Oryctes monoceros, Cirina
forda, Nsike, Kigelegele, Kansenda, Bangwangwa, Maguina, Mingungu, Ngohangoha, Bikolongo and Bachache
were inventoried as a source of food in Fizi, Kabare, Mwenga and Walungu Territories (Table 2, Fig. 2), belong-
ing to nine families including Termitidae, Gryllidae, Curculionidae, Tenebrionidae, Acrididae, Gryllotalpidae,
Apidae, Saturniidae and Scarabaeidae and five orders including Isoptera, Orthoptera, Coleoptera, Hymenoptera
and Lepidoptera.
Among the inventoried edible insects, four (M. subhyalinus, A. domesticus, R. phoenicis and A. diaperi-
nus) were found in Fizi, eleven (M. subhyalinus, A. domesticus, R. differens, G. africana, A. mellifera larvae, N.
septemfasciata, L. migratoria, R. ferrugineus, I. oyemensis, I. epimethea and Nsike) in Kabare, seventeen (M.
subhyalinus, A. domesticus, R. phoenicis, A. diaperinus, R. ferrugineus, I. oyemensis, I. epimethea, O. monoceros,
C. forda, Kigelegele, Kansenda, Bangwangwa, Maguina, Mingungu, Ngohangoha, Bikolongo and Bachache) in
Mwenga, and eight (M. subhyalinus, A. domesticus, R. differens, G. africana, A. mellifera larvae, N. septemfasciata,
L. migratoria, and Nsike) in Walungu.
The M. subhyalinus and A. domesticus were used as sources of food in the whole study area, namely Fizi,
Kabare, Mwenga and Walungu Territories, while R. phoenicis and A. diaperinus are consumed only in Fizi and
Mwenga. In Kabare and Walungu Territories, R. differens, G. africana, A. mellifera larvae, N. septemfasciata, L.
migratoria and Nsike are used as sources of food. Moreover, R. ferrugineus, I. oyemensis and I. epimethea are used
as food sources in Kabare and Mwenga. However, O. monoceros, C. forda, Kigelegele, Kansenda, Bangwangwa,
Maguina, Mingungu, Ngohangoha, Bikolongo and Bachache are only used as sources of food only in Mwenga
Territory.
As for the stage of consumption, some edible insect species were consumed at the larval stage, including R.
phoenicis, A. diaperinus, A. mellifera larvae, R. ferrugineus, I. oyemensis, I. epimethea, O. monoceros, C. forda,
Maguina, Mingungu, Ngohangoha, Bikolongo, Bachache and Kansenda, and others at the adult stage (M. sub-
hyalinus, A. domesticus, R. differens, G. africana, Nsike, Kigelegele, and Bangwangwa). Unlike the other edible
insects, the egg and pupa of A. mellifera are also consumed.
The wide biodiversity of edible insect species revealed in South Kivu depicts the importance of entomophagy
in the region. Our findings largely agree with that of Bomolo et al.31. They reported a list of eleven edible insect
species belonging to four families in Haut-Katanga Province, confirming that the Democratic Republic of Congo
has a high diversity of edible insects, making it one of the most important biological diversity in Africa. This
biodiversity in terms of edible insects in DRC was also confirmed by Raheem et al.24, who reported on traditional
consumption and rearing of edible insects in Africa, Asia and Europe. Similarly, Kelemu et al.16 noted that most
edible insects consumed in DRC belong to the orders reported in our findings.
This diversity could be associated with the richness of the natural environment ­conditions33 in each Terri-
tory as most edible insects are gathered from the w ­ ild34. In addition, the geographic distribution of host plants
influences the availability of certain edible insects. A low number of caterpillar species has been attributed to
marked deforestation, forest degradation and p ­ ollution35,36. This situation will likely worsen with the growing

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 4

Vol:.(1234567890)
www.nature.com/scientificreports/

Territory
Common name Scientific name Family Order Fizi Kabare Mwenga Walungu Local name Stage of consumption
Termite Macrotermes subhyalinus Termitidae Isoptera + + + + Lolongue/Bushungwe Winged adult
Makelele/Njanjala /hun-
House cricket Acheta domesticus Gryllidae Orthoptera + + + + Adult
gwe/Ntoro
Palm weevil Rhyncophorus phoenicis Curculionidae Coleoptera + − + − Ebungu/Sololo/Mpose Larvae
Beetle Oryctes rhinoceros Scarabaeidae Coleoptera + − + − Sungunya/Njukisha Larvae and adult
Grasshopper Ruspolia differens Acrididae Orthoptera − + − + Minunu Adult
Mole cricket Gryllotalpa Africana Gryllotalpidae Orthoptera − + − + Nkwananzi Adult
Honey bee Apis mellifera Apidae Hymenoptera − + − + Magusha/Manyagu Egg, larvae and pupa
Red locust Nomadacris septemfasciata Acrididae Orthoptera − + − + Mundurha Adult
Migratory locust Locusta migratoria Acrididae Orthoptera − + − + Tondé Adult
Red palm weevil Rhynchophorus ferrugineus Curculionidae Coleoptera − + + − Bivumbe Larvae
Caterpillar Imbrasia oyemensis Saturniidae Lepidoptera − + + − Milanga Larvae
Caterpillar Imbrasia epimethea Saturniidae Lepidoptera − + + − Taku/Tukumombo Larvae
Rhinoceros beetle Oryctes monoceros Scarabaeidae Coleoptera − − + − Batumbu Larvae
Misigi Cirina forda Saturniidae Lepidoptera − − + − Misigi Larvae
Kansenda NI Curculionidae Coleoptera − − + − Kansenda Larvae
Nsike Gnathocera trivittata Scarabaeidae Coleoptera − + − + Nsike Adult
Kigelegele NI NI NI − − + − Kigelegele Adult
Bangwangwa NI NI NI − − + − Bangwangwa Adult
Maguina NI NI NI − − + − Maguina Larvae
Mingungu NI NI NI − − + − Mingungu Larvae
Ngohangoha NI NI NI − − + − Ngohangoha Larvae
Bikolongo NI NI NI − − + − Bikolongo Larvae
Bachache NI NI NI − − + − Bachache Larvae

Table 2.  Commonly consumed edible insects in selected Territories of South-Kivu. + Insects present and
consumed, − Insects not present, NI not identified. All of these edible insects are identified by local names,
mainly in the dialects Kibembe (Fizi), Kirega (Mwenga) and Mashi (Kabare and Walungu) that are attached to
specific physical characteristics or uses.

human populations and declining forest b ­ ase37. Previous studies reported that edible insect’s consumption and
preference are also influenced by their availability and c­ ultures38,39.
Findings from this study line with Raheem et al.24, who reported that more than a thousand insect species
are worldwide consumed at some stage of their life cycle. In addition, Lepidoptera consumed as caterpillars and
Hymenoptera are mostly eaten in their larval or pupal stages like the A. mellifera mentioned from the survey to
be consumed as egg and pupa. Adults and larvae of Coleoptera are consumed, while the Orthoptera, Isoptera
and Hemiptera orders are mostly consumed as mature a­ dults16. Moreover, Kulma et al.40 investigated the effect of
developmental stage on the nutritional value of edible insects (Blaberus craniifer and Zophobas morio) reported
no significant variations in basic nutrient content in protein quality expressed as the essential amino acid index.
In contrast, they reported a significant difference in protein digestibility, fat content and lipid quality.

Consumer preference for edible insects.  The inventoried edible insects were appreciated differently
(Fig. 3). In Fizi, the most preferred edible insects were the R. phoenicis (41%), A. diaperinus (26%), M. subhyali-
nus (20%), and A. domesticus (13%), while in Kabare, the most preferred were R. differens (55%) and A. mellifera
larvae (45%). Furthermore, I. oyemensis (65%), I. epimethea (20%), R. ferrugineus (11%) and R. phoenicis (4%)
were the most preferred in Mwenga, whileR. differens (39%), M. subhyalinus (22%), G. africana (21%), Nsike
(10%), N. septemfasciata (7%) and A. mellifera larvae (1%) were most preferred in Walungu. Some edible insects
were preferred in more than one Territory, such as M. subhyalinus (Fizi and Walungu), R. phoenicis (Fizi and
Mwenga), R. differens and A. mellifera larvae (Kabare and Walungu).
Familiarity appears to be the key driving force, allowing most respondents to react positively to all edible
species in terms of their willingness to eat them and thus address food and nutrition insecurity and related
issues. This preference has been reported to be linked to a­ vailability13,41, ethnicity/cultures42, ­palatability14 and
­seasonality43, as some species are more prevalent and familiar in some Territories than others. In addition, indig-
enous knowledge and processing can also influence the preference of edible insect s­ pecies41.
Bomolo et al.31 revealed that caterpillar consumption is common among ethnic populations in D.R. Congo.
This is not the case for Walungu, Kabare and Fizi, although it is the most appreciated edible insects in Mwenga.
In Katanga, the Bemba and Lamba tribes have a long history of caterpillar consumption since the eighteenth
and nineteenth ­centuries44, while the other tribes (Musanga, Tetela, Mongo, Baluba, Rund, Bacongo, Katshokwe,
Emba, Songe, Ndembo, Kaminungu, Kalwena, Kete, Basankusu, Kanyoka, Sanga, Mbote, Yombe, etc.) appear
to lack a strong history of caterpillar consumption. In addition, some of these tribes are reluctant to engage in

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 5

Vol.:(0123456789)
 www.nature.com/scientificreports/

Figure 2.  (a) Macrotermes subhyalinus (Termite); (b) Acheta domesticus (House cricket); (c) Rhyncophorus
phoenicis larvae (Palm weevil larvae); (d) Alphitobius diaperinus larvae (Beetle); (e) Ruspolia differens
(Grasshopper); (f) Apis mellifera larvae (Honey bee); (g) Locusta migratoria (Migratory locust); (h)
Rhynchophorus ferrugineus larvae (Red palm weevil); (i) Imbrasia oyemensis (Caterpillar); (j) Oryctes monoceros
larvae (Rhinoceros beetle); (k) Imbrasia epimethea (Caterpillar); (l) Gryllotalpa Africana (Mole cricket); (m)
Gnathocera trivittata (Nsike); (n) Nomadacris septemfasciata (Red locust; (o) Cirina forda (Misigi). (Images
mixed using Microsoft PowerPoint for Mac 16.29.1 https://​www.​micro​soft.​com).

entomophagy, specifically to consume caterpillars due to their religious ­beliefs45. Studies have shown that educa-
tion would play a crucial role in increasing the positive attitude towards edible insects among c­ onsumers13,46.

Preference for edible insects.  The plotted data (Fig. 4) shows the appreciation factors for the most pre-
ferred inventoried edible insects (G. africana, R. differens, A. mellifera larvae, Nsike, N. septemfasciata and I.
oyemensis) represented in two Territories (n = 260). Respondents rated them according to taste, size, shape,
nutritional value and colour. Most of these insects were valued for their taste, especially R. differens (33%), I.
oyemensis (32%), A. mellifera larvae (17%), G. africana (3%), and Nsike (2%), except for N. septemfasciata, which
was valued for its size (3%) and shape (0.4%). As for size, R. differens (5%), G. africana (4%) and N. septemfas-
ciata (3%) were the most valued. Most of these edible insects were not valued for their nutritional value or colour
except for A. mellifera larvae (0.4%) and I. oyemensis (1%) for nutritional value and R. differens (3%) and Nsike
(1%) for colour.
Considering M. subhyalinus and A. domesticus represented in all Territories (n = 520). Irrespective of Ter-
ritories, their appreciation depended only on their taste, size and shape (Fig. 5). More appreciation was based
on the taste with 8% and 3% against 2% and 1% for the size for M. subhyalinus and A. domesticus, respectively;
only M. subhyalinus was appreciated for its shape (1%).
Entomophagy habits differ from country to country and culture to culture, as do preference factors. Insect
consumption depends not only on sensory c­ haracteristics13 and nutritional ­value10,41 but also on customs, ethnic
preferences, ­prohibitions7, and medicinal p ­ roperties47. Insects were once associated with filth, fear of contami-
nation and disease, as well as psychological and biased thinking about taste, smell, and c­ olour48, with a sense of
disgust that entomophagy was motivated by starvation and is merely a survival ­mechanism13. This is far from
the truth, as insects are not inferior to other protein sources, such as fish, chicken, and beef. However, it will
take a bit more motivation to reverse this ­mentality49. It is possible to explore edible insects for consumption
and increase the possibility of replacing animal products with insects, given that there is evidence that they are
clean, tasty, and ­nutritious50.
In addition, insects have too many ecological advantages over other animal protein ­sources6. Some studies in
European countries such as the N ­ etherlands51 on the acceptance of entomophagy have shown that people who
have eaten insects in the past show significantly more positive attitudes towards entomophagy than people who
have not and are more likely to eat them again. Therefore, it seems important to encourage people to take the
"first step" and familiarize them with insect consumption. Therefore, consumer "education" about entomophagy
should be practiced in its broadest s­ ense51.

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 6

Vol:.(1234567890)
www.nature.com/scientificreports/

Fizi
Kabare
Termite
50 Termite
40 60
Nsike 50 House cricket
30 40
20 Imbrasia epimethea 30 Grasshopper
20
10 10
Beetle 0 House cricket 0
Imbrasia oyemensis Mol e cricket

Red palm weevil Honey bee

Migratory l ocust Red locust

Palm weevil

Termite Termite
Bachache 70 House cricket 40
60
Bikolongo Palm weevil Nsike 30 House cricket
50
40
Ngohangoha Beetle 20
30
20 10
Mingungu 10 Red palm weevil
0 Migratory l ocust 0 Grasshopper

Maguina Imbrasia oyemensis

Bangwangwa Imbrasia epimethea

Red locust Mol e cricket
Kansenda Rhinocer os beetle
Mwenga Kigel egele Misigi
Honey bee Walungu
Figure 3.  Most preferred edible insects in each Territory (n = 130). This is the number of times each edible
insect is preferred. The preference is expressed in percentage. (Figures plotted using Microsoft Excel for Mac
16.56 and mixed using Microsoft PowerPoint for Mac 16.29.1 https://​www.​micro​soft.​com).

35.00 Taste
Factors of preference (%)

30.00 Size
Shape
25.00
Nutritional value
20.00
Color
15.00
10.00
5.00
0.00
t

er

st

sis

tle
l
ke

vi
be

sik

he

vi
cu
pp

e
en

ee
ic

ee
et

Be
N

lo
y
ho
cr

w
m

im
e

w
on

d
ss

ye

lm
e

Re

ep

m
ol

ra

l
pa
M

Pa
G

ia
ia

as

d
as

Re
br
br

Im
Im

Figure 4.  Preference of edible insects represented in two Territories. Respondents gave reasons for preferring
one edible insect over another. The preference was based on taste, size, shape, nutritional value and colour.
(Figures plotted using Microsoft Excel for Mac 16.56 https://​www.​micro​soft.​com).

Seasonal availability of various inventoried edible insects.  Regardless of Territory, three edible
insect groups, mainly A. domesticus, A. diaperinus and A. mellifera larvae, are abundant throughout the year
(Table 3). Nomadacris septemfasciata is also available throughout the year but abundant only during 5 months
and less abundant from February to August. Gryllotalpa africana and R. ferrugineus, on the other hand, were
abundant throughout the rainy season and are less abundant in the dry season. G. africana and R. ferrugineus are
not abundant, respectively, in August and June. Both species are all unavailable in July. Elsewhere, M. subhyali-

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 7

Vol.:(0123456789)
 www.nature.com/scientificreports/

Termite House cricket

Factors of preference (%)

9.00
8.00
7.00
6.00
5.00
4.00
3.00
2.00
1.00
0.00
Taste Size Shape Nutritional value Color

Figure 5.  Preference of Termites and House crickets in the study area. Respondents gave reasons for preferring
one edible insect over another. The preference was based on taste, size, shape, nutritional value and colour.
(Figures were plotted using Microsoft Excel for Mac 16.56 https://​www.​micro​soft.​com).

Rain season Dry season Rain season

Insect species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec TMA TLA
Termite + + + + − 0 0 0 − − + + 6 3
House cricket + + + + + + + + + + + + 12 0
Palm weevil 0 0 0 + + + + − 0 0 + + 6 1
Beetle + + + + + + + + + + + + 12 0
Grasshopper + − − + + 0 0 0 − + + + 6 3
Mole cricket + + + + + 0 0 − + + + + 9 1
Honey bee + + + + + + + + + + + + 12 0
Red locust + − − − − − − − + + + + 5 7
Migratory locust − − 0 0 0 0 0 0 0 0 + + 2 2
Red palm weevil + + + + + − 0 0 + + + + 9 1
Imbrasia oyemensis 0 + + + − 0 0 0 0 0 0 0 3 1
Imbrasia epimethea 0 + + − 0 0 0 0 0 0 0 0 2 1
Rhinoceros beetle + + + − − 0 0 0 0 0 0 + 4 2
Misigi + + + − 0 0 0 0 0 0 0 + 4 1
Kansenda 0 0 + − 0 0 0 0 0 0 0 0 1 1
Nsike 0 0 0 + + + − − 0 0 0 0 3 2
Kigelegele 0 + − 0 0 0 0 0 0 0 0 0 1 1
Bangwangwa − 0 0 0 0 0 0 0 0 0 0 + 1 1
Maguina 0 + + − 0 0 0 0 0 0 0 0 2 1
Mingungu 0 0 + + + − 0 0 0 0 0 0 3 1
Ngohangoha 0 0 0 0 0 0 + + + + − 0 4 1
Bikolongo 0 0 + + + − 0 0 0 0 0 0 3 1
Bachache 0 + + + − 0 0 0 0 0 0 0 3 1

Table 3.  Seasonal availability of various inventoried edible insects. + month of availability, − month of less
availability, 0 month of none availability, TMA Total month of availability, TLA Total month of less availability.

nus, R. phoenicis and R. differens are available half the year (6 months). However, M. subhyalinus and R. differens
were less available for 3 months of the year while R. phoenicis are only available for one month. Other groups
such as Oryctes monoceros, Cirina forda, Ngohangoha, I. oyemensis, Nsike, Mingungu, Bikolongo and Bachache
were abundant for only 3 to 4 months of the year. In contrast, I. epimethea, Maguina, Kigelegele, Kansenda and
Bangwangwa were the most available and abundant for only one to two months of the year.
Availability is an important factor in the consumption of edible ­insects14. Our results corroborate of those of
Ebenebe et al.20, who pointed out that most of the harvesting is done during the rainy season in Nigeria, especially
for winged termites, cricket, caterpillars, A. domesticus, G. africana, and greenish beetle. Smith and P ­ aucar52
suggested that vibrations caused by rain and the sound of thunder would trigger their emergence. Chakravorty
et al.14 also confirmed that the availability of edible insects is seasonal, stating that peak numbers of edible beetles
occur from June to September before decreasing in winter and early spring. They also reported that Odonata
and Orthoptera were most abundant in September and October (late summer).
Insects of the order Hemiptera and Hymenoptera are less abundant from November to February (winter),
while others such as bugs and ants are available throughout the year. This would be attributed to seasonal

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 8

Vol:.(1234567890)
www.nature.com/scientificreports/

Host plants
Insect species Common name Scientific name Signs of presence
Termite NA NA NA
House cricket NA NA NA
Palm weevil Palm, coconut and African oil palm Raffia palm, Cocos nucifera and Elaeis guineensis Cracking noises in palm trunks and odour
Raffia palm, Peltophorum pterocarpum and Man- Indication of its activity at the hole of entrance and
Beetle Palm, yellow flame and mango
gifera spp cracking noises in the palm
Digitaria sp, Panicum maximum, and Sporobolus
Grasshopper Grass, guinea grass and giant rat’s tail grass NA
pyramidalis
Mole cricket NA NA Whistling and canals in the wet ground
Honey bee NA NA NA
Maize, rice, soybean, sugarcane, groundnut and Zea mays, Oryza sativa, Glycine max, Saccharum
Red locust Whistling
sweet potato officinarum, Arachis hypogaea and Ipomoea batatas
Maize, rice, soybean, sugarcane, groundnut and Zea mays, Oryza sativa, Glycine max, Saccharum
Migratory locust Whistling
sweet potato officinarum, Arachis hypogaea and Ipomoea batatas
Cocos nucifera, Elaeis guineensis and Saccharum Indication of its activity at the entrance hole, odour
Red palm weevil Coconut, African oil palm and sugarcane
officinarum, and cracking noises in the palm
Imbrasia oyemensis Red mangrove Rhizophora mangle Caterpillar smells and typical bird songs
Imbrasia epimethea Red mangrove and African blackwood Rhizophora mangle and Erythrophleum africanum Caterpillar smells and typical bird songs
Indication of its activity at the hole of entrance and
Rhinoceros beetle Coconut and African oil palm Cocos nucifera and Elaeis guineensis
Cracking noises in the palm
Misigi Red mangrove and African blackwood Rhizophora mangle and Erythrophleum africanum Whistling
Kansenda Palm and coconut Raffia palm and Cocos nucifera Cracking noises in palm trunks and odour
Jaragua grass, Weeping lovegrass and Giant rat’s Hyparrhenia rufa, Eragrostis curvula and Sporobolus
Nsike NA
tail grass pyramidalis
Kigelegele NA NA NA
Bangwangwa NA NA NA
Maguina NYD NYD NYD
Mingungu NYD NYD NYD
Ngohangoha NYD NYD NYD
Bikolongo NYD NYD NYD
Bachache NYD NYD NYD

Table 4.  Host plants of various consumed edible insects. NA Not applicable, NYD Not yet determined.

changes in various regions of the world. In the Central African Republic, the average consumption of caterpil-
lars increases due to their greater abundance during the rainy ­season13, as seasonal availability and edible insect
consumption are c­ orrelated15. In contrast to our findings, all developmental stages of R. differens can be found
throughout the year in non-swarming populations but at low densities during dry seasons and high densities
during rainy s­ easons53.

Host plants for various inventoried edible insects.  Some edible insects such as M. subhyalinus, A.
domesticus, G. africana, A. mellifera larvae, Kigelegele and Bangwangwa do not necessarily have host plants.
However, R. phoenicis, A. diaperinus, R. differens, N. septemfasciata, L. migratoria, R. ferrugineus, I. oyemensis,
I. epimethea, O. monoceros, C. forda, Kansenda and Nsike require host plants to serve either for habitat or food
source (Table 4). Edible insects such as N. septemfasciata and L. migratoria are dangerous as they use maize (Zea
mays), rice (Oryza sativa), soybean (Glycine max), sugar cane (Saccharum officinarum), groundnut (Arachis
hypogaea) and sweet potato (Ipomoea batatas) crops as host plants, while the latter are also sources of staple
foods for humans. On the other hand, other species R. phoenicis, A. diaperinus, R. ferrugineus, O. monoceros and
Kansenda, were hosted on Raffia palm, Cocos nucifera, Elaeis guineensis, and Mangifera spp trees which are not
only sources of food for humans and a source of income for many people. Their signs of presence differ from one
to another. The presence of R. phoenicis, A. diaperinus, R. ferrugineus, O. monoceros and Kansenda is noticed by
cracking noises in palm trunks, odour, and their activities at the hole of the entrance. Furthermore, caterpillar
smells and typical bird songs were signs of presence for I. oyemensis and I. epimethea. Moreover, G. africana, N.
septemfasciata, L. migratoria and C. forda are noticed by whistling and canals in the wet ground.
Ebenebe and c­ ollaborator20 highlighted that certain edible insects are associated with the following host plants:
cricket-yam; yam beetle-yam; African palm weevil-raffia palm; Rhinoceros beetle-raffia palm, oil palm, coconut
tree; butterfly-iroko (Chlorophora excelsa), locust bean seed (Parkia biglobosa), flamboyant tree (Delonix regia),
croton (Croton tiglium) and ngwu tree; grasshopper and honey bee-Jatropha gossyplifolia, Citrus sinensis, Morinda
lucida, Psidium guajava and Sarcocepha laifolius. According to Ngute et al.54, five of the eleven caterpillar species
studied in central Cameroon were reported to have only one host plant, while others had more than one. They
identified eighteen plants, of which eleven are restricted to natural forest habitats, including Entandrophragma
cylindricum and Baillonella toxisperma. Although many of the identified caterpillar host plants are generally

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 9

Vol.:(0123456789)
 www.nature.com/scientificreports/

Insect species Harvesting techniques Harvesting period Processing methods Preservation techniques
Light trapping near a container.
Termites attracted by light fall inside De-winged, roasted or dry-fried; also
Termite During and after the first rains Drying
before being collected and have their eaten raw
wing removed
House cricket Trapping and handpicking Any time De-winged, roasted or dry-fried Drying
Handpicking after signs of their pres- Gut removed, boiled, fried or roasted,
Palm weevil Any time based on signs Drying
ence is detected sometimes prepared in stews
Handpicking after signs of their pres- Washed, boiled, fried or roasted,
Beetle Any time based on signs Drying
ence is detected sometimes prepared in stews
During the swarming season, the light Soon after the dark for light trapping
trapping technique is used to attract and morning before the sun rises Dry-fried during the swarming
Grasshopper De-winged, roasted or dry-fried
grasshoppers in addition to handpick- when they could not fly for handpick- season
ing on host plants ing
In the evening, following their small De-winged, roasted or dry-fried and
Mole cricket Handpicking Drying
holes boiled
Collecting honeycomb from the hive
Honey bee At night preferably Boiled None
following with honey extraction
Red locust Handpicking on host plants Morning time De-winged, roasted or dry-fried Drying
Migratory locust Handpicking on host plants Morning time De-winged, roasted or dry-fried Drying when it is enough
Handpicking after signs of their pres- Washed, boiled, fried or roasted,
Red palm weevil Any time based on signs Drying
ence is detected sometimes prepared in stews
Handpicking: caterpillar directly
Any time, preferably morning and Boiled, fried or roasted, sometimes
Caterpillar picked after signs of their presence are Drying
evening hours prepared in stews
detected
Handpicking after signs of their pres- Any time, preferably morning and Boiled, fried or roasted, sometimes
Rhinoceros beetle Drying
ence are detected evening hours prepared in stews
Handpicking after signs of their pres- Any time, preferably morning and Boiled, fried or roasted, sometimes
Misigi Drying
ence are detected evening hours prepared in stews
Handpicking after signs of their pres- Any time, preferably morning and
Kansenda Fried Drying
ence are detected evening hours
During the sunny period, they are easy
Dry-fried during the swarming
Nsike Handpicking on host plants to identify and handpicking on the top De-winged, roasted or dry-fried
season
of the host plants
Handpicking after signs of their pres- Any time, preferably morning and
Kigelegele Boiled and Fried Drying
ence are detected evening hours
Handpicking after signs of their pres- Any time, preferably morning and
Bangwangwa Boiled and roasted Drying
ence are detected evening hours
Handpicking after signs of their pres- Any time, preferably morning and Boiled, fried or roasted, sometimes
Maguina Drying
ence are detected evening hours prepared in stews
Handpicking after signs of their pres- Any time, preferably morning and
Mingungu Boiled and roasted Drying
ence are detected evening hours
Handpicking after signs of their pres- Any time, preferably morning and Boiled, fried or roasted, sometimes
Ngohangoha Drying
ence are detected evening hours prepared in stews
Handpicking after signs of their pres- Any time, preferably morning and
Bikolongo Boiled and roasted Drying
ence are detected evening hours
Handpicking after signs of their pres- Any time, preferably morning and Boiled, fried or roasted, sometimes
Bachache Drying
ence are detected evening hours prepared in stews

Table 5.  Harvesting and processing techniques for various consumed edible insects in selected Territories.

wild, a few are domesticated and grown in home gardens and agroforestry systems such as Mangifera indica and
Dacryodes edulis, or are in the process of domestication such as Ricinodendron heudelotii, B. toxisperma and E.
cylindricum54. Also, it should be noted that most of the hosts are plants used as a source of food and revenue;
for example, B. toxisperma is a class A timber species, which produces fruits with a highly valued and edible oil
is ­extracted55.
It has been reported that out of 21,252 observations, R. differens were observed 20,915 (98%) times on grasses
and sedges, with a total of 19 grass species (Poaceae) and two sedge species (Cyperaceae). Among the grasses the
dominant species were P. maximum, B. ruziziensis, C. gayana, H. rufa, Cynodon dactylon, Sporobolus pyramidalis
and P. purpureum53. When reared in the laboratory, R. differens accept artificial food, leaves, flowers, and grains
of many types of grass, including cultivated c­ ereals56,57. At this point, our findings line with those of Meutchieye
et al.19, who observed that the cracking noises in the palm trunks three to four weeks after the final collection of
palm wine, the yellow of the internal raffia bamboo, caterpillar odours and typical bird songs and whistling are
the signs indicating the presence of R. phoenicis, caterpillars, R. differens and field crickets.

Harvesting and processing techniques of edible insects.  Thistudyfound that harvesting time and
techniques vary according to local knowledge, practices, and insect species (Table 5). Three harvesting tech-
niques were identified, including trapping, collection. M. subhyalinus (during and after the first rains) and R.

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 10

Vol:.(1234567890)
www.nature.com/scientificreports/

differens (during the swarming season) are trapped with light near a container; once attracted by light, they fall
inside before being collected then de-winged. In addition, R. differens are also hand-picked on host plants while
A. domesticus are collected by trapping and handpicking at any time. Furthermore, R. phoenicis, A. diaperinus, G.
africana, A. mellifera larvae, N. septemfasciata, L. migratoria, R. ferrugineus, I. oyemensis, I. epimethea, O. monoc-
eros, C. forda, Nsike, Maguina, Mingungu, Ngohangoha, Bikolongo and Bachache are hand-picked.
Some edible insects are collected only in the morning or evening, while others are collected at any time. The
harvest decreases with the intensity of the sun. Ruspolia differens are light trapped soon after the dark. Like N.
septemfasciata, L. migratoria and Nsike, R. differens are also hand-picked in the morning before the sun rises when
they cannot fly. On the other hand, A. mellifera larvae and G. africana are collected at night and in the evening,
preferably. Rhyncophorus phoenicis, A. diaperinus, R. ferrugineus, I. oyemensis, I. epimethea, O. monoceros, C.
forda, Maguina, Mingungu, Ngohangoha, Bikolongo and Bachache are collected Any time during the day, but
preferably in the morning and evening hours.
Processing methods and preservation techniques vary according to the type of edible insect and the purpose,
whether for direct consumption or preservation. Generally, they are all dried or dry-fried for preservation except
for the honeybee (Table 5). Most edible insects with wings such as M. subhyalinus, A. domesticus, R. differens,
G. africana, N. septemfasciata, L. migratoria and Nsike are de-winged before being roasted, dry-fried, fried or
boiled for G. africana. M. subhyalinus are also eaten raw. Rhyncophorus phoenicis, A. diaperinus, R. ferrugineus, I.
oyemensis, I. epimethea, O. monoceros, C. forda, Maguina, Ngohangoha and Bachache are gut removed, washed,
boiled, fried or roasted, and sometimes prepared in stews. A. mellifera larvae are boiled while Kigelegele, Bang-
wangwa, Mingungu, and Bikolongo are boiled and roasted.
These results are consistent with those of Meutchieye et al.19, as well as those of Ebenebe et al.20, who also
pointed out that hand picking is one of the major techniques used to harvest crickets, Rhinoceros beetle, stinking
grasshopper (in the early hours of the morning), African palm weevil (from rotting palm tree) and caterpillars.
In contrast, light trapping is used to harvest termites, green grasshoppers, house cricket and mole cricket. In
addition, Ebenebe et al.20 added that crickets were also harvested by digging out from the tunnels during the dry
season and that house cricket was also harvested by digging out from small sound mounds with depression that
indicates their entrance. In contrast to the results of this study, Ebenebe et al.20 stated that mole crickets were
harvested by light trapping. Singing a "Wee wee" song at the base of the host tree is another notable technique
used to harvest the caterpillar of a particular moth species. The more you sing, the more they fall from the top
of the tree and are harvested.
To harvest enough R. differens by light trapping during the swarming season, locals lit the lamps/bulbs out-
side the houses and secure the areas where these lamps/bulbs are located with corrugated iron sheets bent into
a cone shape leading to baskets, small drums where the falling R. differens are ­harvested58. For non-swarming
R. Differens, harvesting is done early in the morning between 6 and 7 AM, when they are inactive and unable
to ­fly17. Dao et al.59 also reported two harvesting techniques: direct termite collections from their mounds and
termite trapping in containers. The direct collection involves partially destroying termite mounds, while trapping
implies looking for signs of termite presence such as mud sheets and tracks on the ground. Techniques depend
on the termite genus. Small termite mounds of Trinervitermes and Cubitermes can be dug with a hoe or pickaxe,
and the mounds are collected in the morning between 6 and 9 AM. The traps are placed between 6 and 8 AM or
in the evening around 6 PM. They are covered with foliage or pieces of cloth to protect them from the sun. The
same harvesting techniques were recently described in G ­ hana60.
It was reported that edible insects were sun-dried, baked, steamed and processed into crackers, muffins and
sausage ­meat61,62. Our findings confirm those of Ebenebe et al.20, who also found that salted roasting is one of
the techniques used to process termites, crickets, Rhinoceros beetle, grasshopper and locust. They added that
grasshopper and locust are consumed dried as well. On the other hand, African palm weevils are consumed raw
or fried with pepper sauce. This study found that drying was the most used preservation technique as it is the
most widely used technology for increasing the shelf-life of foods. The drying ranges from traditional, including
roasting, frying and solar drying, to modern freeze-drying and microwave-assisted d ­ rying63.

Conclusion and recommendations
The wide biodiversity of edible insects observed depicts the importance of entomophagy in the region. Twenty-
three edible insects were recorded belonging to nine families and five orders. These insects are consumed as
larvae, adults or as eggs and pupa. About four of them were the most popular edible insects in Fizi and Mwenga,
two in Kabare, and six in Walungu Territory. Additionally, R. differens, I. oyemensis, A. mellifera larvae, G. africana
and Nsike were the most preferred for their taste. Besides, some are available throughout the year, while the oth-
ers are less available. Many plants have been recorded as host plants, including plants used as food sources, feed
and income for humans. Harvesting strategies and time, processing methods, and preservation techniques vary
according to local knowledge and practices and insect species. These findings suggest similar studies in other
provinces and further research on the nutritional and safety profiling of processed and non-processed edible
insects while encouraging the rearing of certain edible insects for mass production as the demand is too high.

Received: 23 July 2021; Accepted: 13 January 2022

References
1. United Nations. Publications World Population Prospects 2019: Highlights (Springer, 2019).
2. Godfray, J. F., Pretty, J., Robinson, S., Thomas, S. M. & Toulmin, C. Food security: The challenge of feeding 9 billion people. Science
327, 812–819 (2010).

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 11

Vol.:(0123456789)
 www.nature.com/scientificreports/

3. Premalatha, M., Abbasi, T., Abbasi, T. & Abbasi, S. A. Energy-efficient food production to reduce global warming and ecodegrada-
tion: The use of edible insects. Renew. Sustain. Energy Rev. 15, 4357–4360 (2011).
4. Halloran, A., Vantomme, P., Hanboonsong, Y. & Ekesi, S. Regulating edible insects: The challenge of addressing food security,
nature conservation, and the erosion of traditional food culture. Food Secur. https://​doi.​org/​10.​1007/​s12571-​015-​0463-8 (2015).
5. Kinyuru, J. N., Mogendi, J. B., Riwa, C. A. & Ndung, N. W. Edible insects: A novel source of essential nutrients for human diet:
Learning from traditional knowledge. Anim. Front. 1, 14–19. https://​doi.​org/​10.​2527/​af.​2015-​0014 (2015).
6. Rumpold, B. A. & Schlüter, O. K. Potential and challenges of insects as an innovative source for food and feed production. Innov.
Food Sci. Emerg. Technol. 17, 1–11 (2013).
7. Van Huis, A. & Oonincx, D. G. A. B. The environmental sustainability of insects as food and feed: A review. Agron. Sustain. Dev.
https://​doi.​org/​10.​1007/​s13593-​017-​0452-8 (2017).
8. Kou, L. & Adámková, A. Nutritional and sensory quality of edible insects. NFS J. 4, 22–26 (2016).
9. Mishyna, M. & Chen, J. Sensory attributes of edible insects and insect-based foods: Future outlooks for enhancing consumer
appeal. Trends Food Sci. Technol. 95, 141–148 (2020).
10. Kinyuru, J. N. et al. The role of edible insects in diets and nutrition in East Africa. In Edible Insects in Sustainable Food Systems
(eds Halloran, A. et al.) 93–108 (Springer, 2018).
11. Payne, C. L. R., Scarborough, P., Rayner, M. & Nonaka, K. A systematic review of nutrient composition data available for twelve
commercially available edible insects, and comparison with reference values. Trends Food Sci. Technol. 47, 69–77 (2016).
12. Sánchez-Muros, M. J., Barroso, F. G. & Manzano-Agugliaro, F. Insect meal as renewable source of food for animal feeding: A
review. J. Clean. Prod. 65, 16–27 (2014).
13. Van Huis, A. Potential of insects as food and feed in assuring food security. Annu. Rev. Entomol. https://​doi.​org/​10.​1146/​annur​
ev-​ento-​120811-​153704 (2013).
14. Chakravorty, J., Ghosh, S. & Meyer-rochow, V. B. Comparative survey of entomophagy and entomotherapeutic practices in six
tribes of Eastern Arunachal Pradesh (India). J. Ethnobiol. Ethnomed. 1, 1–12 (2013).
15. Takeda, J. & Sato, H. Multiple subsistence strategies and protein resources of horticulturists in the Zaire basin: The Ngandu and
the Boyela. In Tropical Forests, People and Food: Biocultural Interactions and Applications to Development (eds Hladik, A. et al.)
(Springer, 1993).
16. Kelemu, S. et al. African edible insects for food and feed: inventory, diversity, commonalities and contribution to food security. J.
Insects as Food Feed 1, 103–119 (2015).
17. Kinyuru, J. N., Kenji, G. M., Muhoho, S. N. & Ayieko, M. Nutritional potential of longhorn grasshopper (Ruspolia differens) con-
sumed in Siaya district, Kenya. J. Agric. Sci. Technol. 1, 32–46 (2010).
18. Hanboonsong, Y., Jamjanya, T. & Durst, P. B. Six-legged livestock: Edible insect farming, collection and marketing in Thailand.
RAP Publ. 3, 1–11 (2013).
19. Meutchieye, F., Tsafo, K. E. & Niassy, S. Inventory of edible insects and their harvesting methods in the Cameroon centre region.
J. Insects Food Feed 2, 145–152 (2016).
20. Ebenebe, C. I., Amobi, M. I., Udegbala, C., Ufele, A. N. & Nweze, B. O. Survey of edible insect consumption in south-eastern
Nigeria. J. Insects Food Feed 3, 241–251 (2017).
21. Hartmann, C., Shi, J., Giusto, A. & Siegrist, M. The psychology of eating insects: A cross-cultural comparison between Germany
and China. Food Qual. Prefer. 44, 148–156 (2015).
22. Cicatiello, C., Rosa, B. D., Franco, S. & Lacetera, N. Consumer approach to insects as food: barriers and potential for consumption
in Italy. Br. Food J. 118, 2271–2286 (2016).
23. Laureati, M., Proserpio, C., Jucker, C. & Savoldelli, S. New sustainable protein sources: Consumers’ willingness to adopt insects.
Ital. J. Food Sci. (2016).
24. Raheem, D. et al. Traditional consumption of and rearing edible insects in Africa, Asia and Europe. Crit. Rev. Food Sci. Nutr. 8398,
1–12 (2018).
25. Schlup, Y. & Brunner, T. Prospects for insects as food in Switzerland: A tobit regression. Food Qual. Prefer. https://​doi.​org/​10.​
1016/j.​foodq​ual.​2017.​10.​010 (2018).
26. Wilkinson, K. et al. Australian consumers’ awareness and acceptance of insects as food. Insects https://​doi.​org/​10.​3390/​insec​ts902​
0044 (2018).
27. Woolf, E., Zhu, Y., Emory, K., Zhao, J. & Liu, C. Willingness to consume insect-containing foods: A survey in the United States.
LWT Food Sci. Technol. 102, 100–105 (2019).
28. Doocy, S. et al. Improving household food security in eastern Democratic Republic of the Congo: A comparative analysis of four
interventions. Food Secur. 10, 649–660 (2018).
29. Kim, R., Martin, B., Nielsen, R. & Thorsen, B. J. Household determinants of bushmeat and eru (Gnetum africanum) harvesting for
cash in the Democratic Republic of Congo. Environ. Dev. Sustain. 19, 1425–1443 (2017).
30. Vliet, N. V., Muhindo, J., Nyumu, J. K. & Nasi, R. From the forest to the dish: A comprehensive study of the wildmeat value chain
in Yangambi, Democratic Republic of Congo. Front. Ecol. Evol. 7, 1–10 (2019).
31. Bomolo, O. et al. Ecological diversity of edible insects and their potential contribution to household food security in Haut-Katanga
Province, Democratic Republic of Congo. Afr. J. Ecol. 55, 640–655. https://​doi.​org/​10.​1111/​aje.​12400 (2017).
32. Lrpayne, C., Mato, B. & Fruth, B. Entomophagy in the area surrounding LuiKotale, Salonga National Park, Democratic Republic
of the Congo. Afr. Stud. Monogr. 37, 1–12 (2016).
33. Twine, W., Moshe, D., Netshiluvhi, T. & Siphugu, V. Consumption and direct-use values of savanna bio-resources used by rural
households in Mametja, a semi-arid area of Limpopo province, South Africa. S. Afr. J. Sci. 1, 467–473 (2003).
34. Levang, P., Lescuyer, G., Noumbissi, D. & Déhu, C. Does gathering really pay? Case studies from forest areas of the East and South
regions of Cameroon. For. Trees Livelihoods 24, 128–143 (2015).
35. Ramos-elorduy, J. Threatened edible insects in Hidalgo, Mexico and some measures to preserve them. J. Ethnobiol. Ethnomed. 10,
1–10 (2006).
36. Schabel, H. Forest-based insect industries. In Forest entomology in East Africa: forest insects of Tanzania (2006).
37. Hoare, A. L. The Use of Non-timber Forest Products in the Congo Basin: Constraints and Opportunities (Rainforest Foundation,
2007).
38. Hanboonsong, Y., Jamjanya, T. & Durst, P. B. Six-Legged Livestock: Edible Insect Farming, Collection and Marketing in Thailand
(Springer, 2013).
39. Raubenheimer, D. & Rothman, J. M. Nutritional ecology of entomophagy in humans and other primates. Annu. Rev. Entomol. 58,
141–160. https://​doi.​org/​10.​1146/​annur​ev-​ento-​120710-​100713 (2013).
40. Kulma, M., Kou, L., Homolková, D. & Plachý, V. Effect of developmental stage on the nutritional value of edible insects: A case
study with Blaberus craniifer and Zophobas morio. J. Food Compos. Anal. 92, 103570 (2020).
41. Obopile, M. & Seeletso, T. G. Eat or not eat an analysis of the status of entomophagy in Botswana. Food Secur. 5, 817–824. https://​
doi.​org/​10.​1007/​s12571-​013-​0310-8 (2013).
42. Riggi, L. G., Veronesi, M. & Goergen, G. Observations of entomophagy across Benin: Practices and potentials. Food Secur. 8,
139–149. https://​doi.​org/​10.​1007/​s12571-​015-​0509-y (2016).
43. Kinyuru, J. N. et al. Nutrient composition of four species of winged termites consumed in western Kenya. J. Food Compos. Anal.
30, 120–124 (2013).

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 12

Vol:.(1234567890)
www.nature.com/scientificreports/

44. Malaisse, F. & Latham, P. Human consumption of Lepidoptera in Africa: An updated chronological list of references (370 quoted!)
with their ethnozoological analysis. Geol. Ecol. Trop. Rev. Int. 1, 339–372 (2014).
45. Meyer-rochow, V. B. Food taboos: Their origins and purposes. J. Ethnobiol. Ethnomed. 10, 1–10 (2009).
46. Looy, H. & Wood, J. R. Attitudes toward invertebrates: Are educational “bug banquets” effective?. J. Environ. Educ. 37, 37–48
(2006).
47. Musundire, R., Zvidzai, C. J. & Chidewe, C. Bio-active compounds composition in edible stinkbugs consumed in south-bio-active
compounds composition in edible stinkbugs consumed in South-Eastern Districts of Zimbabwe. Int. J. Biol. https://​doi.​org/​10.​
5539/​ijb.​v6n3p​36 (2014).
48. Deroy, O., Reade, B. & Spence, C. The insectivore’s dilemma, and how to take the West out of it. Food Qual. Prefer. 44, 44–55 (2015).
49. Pliner, P. & Salvy, S. Food neophobia in humans. Front. Nutr. Sci. 3, 75 (2006).
50. Gahukar, R. T. Entomophagy and human food security. Int. J. Trop. Insect Sci. 31, 129–144 (2011).
51. Lensvelt, E. J. S. & Steenbekkers, L. P. A. Exploring consumer acceptance of entomophagy: A survey and experiment in Australia
and the Netherlands. Ecol. Food Nutr. 53, 543–561 (2014).
52. Smith, A. B. T. & Paucar, A. Taxonomic review of Platycoelia lutescens (Scarabaeidae: Rutelinae: Anoplognathini) and a description
of its use as food by the people of the Ecuadorian highlands. Ann. Entomol. Soc. Am. 93, 408–414 (2000).
53. Opoke, R. et al. Host plants of the non-swarming edible bush cricket Ruspolia differens. Ecol. Evol. 9, 3899–3909. https://​doi.​org/​
10.​1002/​ece3.​5016 (2019).
54. Ngute, A. S. K. et al. Edible caterpillars in central Cameroon: Host plants, value, harvesting, and availability. For. Trees Livelihoods
29, 16–33 (2020).
55. Meunier, Q., Moumbogou, C. & Doucet, J.-L. Les arbres utiles du Gabon. (Presses agronomiques de Gembloux, 2015).
56. Malinga, G. M. et al. Mixed artificial diets enhance the developmental and reproductive performance of the edible grasshopper,
Ruspolia differens (Orthoptera: Tettigoniidae). Appl. Entomol. Zool. 53, 237–242 (2018).
57. Valtonen, A. et al. The edible katydid Ruspolia differens is a selective feeder on the inflorescences and leaves of grass species.
Entomol. Exp. Appl. 166, 592–602. https://​doi.​org/​10.​1111/​eea.​12707 (2018).
58. Mmari, M. W., Kinyuru, J. N., Laswai, H. S. & Okoth, J. K. Traditions, beliefs and indigenous technologies in connection with the
edible longhorn grasshopper Ruspolia differens (Serville 1838) in Tanzania. J. Ethnobiol. Ethnomed. 13, 1–11. https://​doi.​org/​10.​
1186/​s13002-​017-​0191-6 (2017).
59. Dao, A. N. C. et al. Traditional methods of harvesting termites used as poultry feed in Burkina Faso. Int. J. Trop. Insect Sci. 40,
109–118 (2020).
60. Boafo, H. A. et al. Use of termites by farmers as poultry feed in Ghana. Insects 10, 1–13. https://​doi.​org/​10.​3390/​insec​ts100​30069
(2019).
61. Ayieko, M., Oriaro, V. & Nyambuga, I. Processed products of termites and lake flies: Improving entomophagy for food security
within the Lake Victoria region. Afr. J. Food Agric. Nutr. Dev. 10, 2085–2098 (2010).
62. Chung, A. Y. C. Edible Insects and Entomophagy in Borneo (Springer, 2010).
63. Melgar-lalanne, G., Hernandez-Alvarez, A.-J. & Salinas-Castro, A. Edible insects processing: Traditional and innovative technolo-
gies. Compr. Rev. Food Sci. Food Saf. 18, 1166–1191 (2019).

Acknowledgements
The authors would like to thank Benjamin Kalalizi, Belanila Victoire, Rebecca Kajuru, Sandra Lukausa and
Tresor Buhendwa for their contribution to data collection. We also extend our acknowledgements to the Uni-
versité Evangélique en Afrique (UEA) and the Regional Universities Forum for Capacity Building in Agriculture
(RUFORUM) for their support (Grant ID: Grant#RU/2020/GTA/DRG/015).

Author contributions
Conceptualization; J.I. and R.A. Methodology; J.I., R.A., J.K., K.K., D.B., S.N. and P.U. Validation; All authors;
Data collection; J.I. Data curation; J.I. and R.A. Formal analysis & drafting paper; J.I. Review & Editing of the
manuscript; All authors.

Competing interests 
The authors declare no competing interests.

Additional information
Correspondence and requests for materials should be addressed to J.I.
Reprints and permissions information is available at www.nature.com/reprints.
Publisher’s note  Springer Nature remains neutral with regard to jurisdictional claims in published maps and
institutional affiliations.
Open Access  This article is licensed under a Creative Commons Attribution 4.0 International
License, which permits use, sharing, adaptation, distribution and reproduction in any medium or
format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the
Creative Commons licence, and indicate if changes were made. The images or other third party material in this
article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the
material. If material is not included in the article’s Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from
the copyright holder. To view a copy of this licence, visit http://​creat​iveco​mmons.​org/​licen​ses/​by/4.​0/.

© The Author(s) 2022

Scientific Reports | (2022) 12:1576 | https://doi.org/10.1038/s41598-022-05607-y 13

Vol.:(0123456789)