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Research Article | | Peer-Reviewed

Diversity, Abundance and the Community Structure of the Flower-Visiting Insects on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) in Bilone (Obala-Cameroon)

Auguste Pharaon Mbianda,  Moukhtar Mohammadou,  Taïmanga,  Andrea Sarah Kenne,  Sedrick Junior Tsekane,  Alice Virginie Tchiaze Ifoue,  Xavier Arthur Nyoumi Ongolo,  Dounia Dounia,  Nadine Esther Otiobo Atibita,  Chantal Douka,  Joseph Blaise Pando,  Fernand-Nestor Tchuenguem Fohouo,  Martin Kenne*
Published in American Journal of Entomology (Volume 9, Issue 1)
Received: 12 December 2024     Accepted: 24 December 2024     Published: 21 January 2025
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Abstract

In order to identify flower-visiting insects on sesame plants and characterize the community structure, ecological survey was conducted in Bilone agroecological farm in 2022 and 2023, in 15 experimental plots (6x5.5 m each) each year, created in a 1,600 m² area. Insects were captured, stored in papillotes (Lepidoptera) or in vials containing 70° alcohol (other adults) and identified at the species level in laboratory. A total of 1,703 specimens were captured. They belonged to five orders, 12 families, 18 genera and 19 species. Hymenoptera was mostly collected order (91.5%) followed by Diptera (4.5%), Lepidoptera (1.8%), Neuroptera (0.9%) and Orthoptera (1.3%). Apidae was the most collected family (42.4%) followed by Formicidae (34.1%), Megachilidae (11.6%) while other families were rare: Acrididae (1.3%), Ascalapidae (0.9%), Calliphoridae (0.5%), Eumenidae (0.7%), Halictidae (2.2%), Muscidae (4.0%), Nymphalidae (1.3%), Pieridae (0.5%), and Vespidae (0.6%). Apis mellifera adansonii (Apidae: 30.6%) was the most recorded species, followed by Paratrechina longicornis (Formicidae: 12.3%), Pheidole megacephala (Formicidae: 9.4%), Myrmicanioa opaciventris (Formicidae: 8.9%), Megachile cincta (Megachilidae: 7.0%), Amegilla calens (Apidae: 6.2%), Xylocopa olivacea (Apidae: 5.6%), Megachile kamerunensis (Megachilidae: 4.6%), Musca domestica (Diptera: 4.0%), Camponotus maculatus (Formicidae: 3.65%), Lasioglossum hancocki (Halictidae: 2.2%), and Pteropera carnapi (Acrididae: 1.3%). Calliphora vicina (Calliphoridae) was recorded exclusively in 2022. Two exotic Diptera (Cl. vicina and Mu. domerstica) were myiasigenic species. The exotic Eumenidae Delta sp. and the afrotropical predator Ascalaphus africanus (Ascalapidae) were recorded as well as the phytophagous Acrididae Pe. carnapi. Potential pests (Nymphalidae, Pieridae and Acrididae) cumulatively represented 3.1% of the collection. The community was highly diversed and lowly dominated by a few species. Ca. maculatus was simply abundant in 2023. Amegilla calens, Ap. mellifera adansonii, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Ph. megacephala and Xy. olivacea were simply abundant. Amegilla calens and Me. cincta, were co-dominants in 2022. Ca. maculatus and Me. kamerunensis were co-dominants in 2023. Apis mellifera adansonii, Pa. longicornis, Ph. megacephala, My. opaciventris and Xy. olivacea were co-dominants in each year. Ca. maculates and Cl. vicina were rare in 2022. Bicyclus dorothea (Nymphalidae), Delta sp. and La. hancocki were rare in 2023. Acraea acerata (Nymphalidae), Ascalaphus africanus (Ascalapidae), Catopsilia florella (Pieridae), Pteropera carnapi (Acrididae) and Synagris conuta (Vespidae) were rare. High value of Motomura constant (m=0.777 in 2022) and Preston constant (m=0.726 in 2023) suggested least evolved pioneer assemblages with species competition limited to the physical space. Overall, flower visiting insects exhibited a global positive net association.

Published in American Journal of Entomology (Volume 9, Issue 1)
DOI 10.11648/j.aje.20250901.14
Page(s) 28-54
Creative Commons

This is an Open Access article, distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution and reproduction in any medium or format, provided the original work is properly cited.

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Copyright © The Author(s), 2025. Published by Science Publishing Group
Previous article
Keywords

Assemblage Composition, Co-Dominant Species, Rare Species, Theoretical Model, Assemblage Functioning, Sesame Plants

1. Introduction
Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) is an annual herbaceous plant and one of the most cultivated oilseed crops. It is native to Asia () and some African countries and was cultivated for over 4,300 years in and
[1, 2]
. It belongs to the division Tracheophyta, class Magnoliopsida, order Scrophulariales or Lamiales, family Pedaliaceae and genus Sesamum
[3]
. According to the Integrated Taxonomic Information System website
[3]
, this genus includes in , three known cultivated valid species: Se. alatum Thonn. native to the dry zone of Africa from Western Sahara to Egypt and south to KwaZulu-Natal in South Africa
[4]
, Se. indicum L. native to India, and Sesamum radiatum Schumach and Thonn, native to the west and central Africa
[5]
. Se. indicum is mainly grown in tropical and subtropical regions of Asia, Africa and with a wide diversity of genotypes
[6-8]
. The plant is erect (0.5- tall in optimal growing conditions, presents green stem, rarely purple, basal diameter: one to three centimeters). Upper leaves are lanceolate, while lower leaves are trilobed (7.5- long). The plant can be glabrous, velvety or hairy, the hairy aspect of the stem and branches being used as grouping factor of varieties
[9]
. The lower leaves are opposite, broad (12 x ), roughly lobed and with a long petiole (about five centimeters) while the upper leaves are alternate or sub-opposite, or narrow (9 x ), with a particular phyllotaxis
[6, 9]
. The leaves are dull green with hairs and stomata on both sides
[6]
. Zygomorphic flowers (bisexual and then hermaphrodite) with pendulous tubular corolla (3- in length) and coloring of various shades of purple white (mostly white or pink) hang down from the stem and they can self-pollinate
[10]
. Flowers occur singly or in groups of two to three in the leaf axils, the androecium consists of four stamens (two long of 1.5-2.0 mm each and two short of 1.0-1.5 mm each) and the gynoecium has superior ovary, multicarpelar and a long style (1.5-2.0 mm) with bifid stigma
[10]
. The flower produces nectar in a nectary disk surrounding the ovary and in a couple of extrafloral nectaries on both sides of the pedicel. Anthesis occurs early in the morning when the stigma becomes receptive and senescence can occur six to 12 hours later, depending on the variety and environmental conditions
[10]
. According the same authors these characteristics of floral biology refer to varieties cultivated especially in warm weather environments, but there is evidence that varieties adapted to tropical conditions behave differently. The genus Sesamum includes many varieties which differ in their dimensions, shape, and type of growth, color of flowers, size, color and composition of seeds
[11]
. The species Se. indicum (synonyms: Se. orientale Sieber ex C. Presl, 1828, Se. edule Steud. (1821), Se. luteum Retz., 1791, Se. oleiferum Moench, 1802, Se. africanum Tod, Se. foetidum Afzel. ex Engl.) is presently cultivated in 65 countries across Asia, Africa, Europe, Central and South America
[11]
. From 2012 to 2016, the global world sesame production was estimated at 12.22 million tons
[12]
and it was estimated in 2021 at 1,150,714 thousand tons, with the yield of 390 kg.ha-1
[10, 13]
. Asia and Africa hold about 90% of the planted area, Egypt, Central Africa, Israel, Peru, Saudi Arabia and Macedonia being the main producing countries of oil crops
[10, 13]
. The sesame production ranks 9th among the 13 main oilseed crops (90% of global edible oil production in the world)
[12, 14]
. Global production of sesame seeds was estimated in 2021 by the Food and Agricultural Organization (FAO) at about 6,667,344 tons grown on 12,965,045 ha with an average yield of about 514 kg.ha-1
[15]
. According to the same source of information, in 2022, it was estimated at about 6,741,479 tons grown on 12,836,776 ha with an average yield of about 525 kg.ha-1. Which showed in 2021 and 2022, a global decrease in cultivated area of about 128,269 ha, an increase in overall production of 74,136 tons and in the yield production of about 11 kg. ha-1. In terms of overall production in 2021, Africa occupied the 1st position (3,997,094 tons grown on 8,417,309 ha) followed in the 2nd position by Asia (about 2,389,914 tons grown on 4,060,409 ha), the 3rd position was occupied by Americas (about 280,295 tons grown on 487,286 ha), and Europe occupied the 5th position (about 41 tons grown on 40 ha)
[15]
. Based on the production yield, in 2021 it was the highest in Europe (about 1,035 kg.ha-1), followed in 2nd position by Asia (about 589 kg.ha-1), in 3rd position by the Americas (about 575 kg.ha-1) and Africa occupied the 4th position (about 475 kg.ha-1)
[15]
. In 2022, the same source of information reported similar estimates (Africa: about 4,000,119 tons grown on about 8,222,425 ha with an average yield of about 487 kg.ha-1; Asia: about 2,401,093 tons grown on 4,010,987 ha with an average yield of about 599 kg.ha-1; Americas: about 340,226 tons grown on 603,324 ha with an average yield of 564 kg.ha-1; Europe: about 41 tons grown on 40 ha)
[15]
. In Africa, comparison of the estimation in 2021 (around four million tons) to the amount produced in the preceding years, showed a decreased number by approximately 633,650 tons
[16]
. The ranking of regions in Africa where sesame is grown showed in 2021 and 2022 that Eastern Africa ranked in the 1st position (2021: 1,436,628 tons grown on 2,380,476 ha with a yield of 604 kg.ha-1; 2022: 1,375,069 tons, 2,363,960 ha and 582 kg/ha), followed in the 2nd position by Northern Africa (2021: 1,285,540 tons, 4,231,280 ha and 304 kg.ha-1; 2022: 1,279,981 tons, 4,184,296 ha and 306 kg.ha-1), in the 3rd position by Western Africa (2021: 814,583 tons, 1,286,685 ha and 633 kg.ha-1; 2022: 874,154.85 tons, 1,167,629 ha and 749 kg.ha-1) and the Central Africa ranked in the 4th position (2021: 460,342 tons, 518,868 ha and 887 kg.ha-1; 2022: 470,914 tons, 506,541 ha and 930 kg.ha-1)
[15]
. The first ten African leading sesame producers in 2021 were Sudan (1,119,026 tons), Tanzania (700,000 tons), Nigeria (440,000 tons), Burkina Fasso (270,000 tons), Tchad (196,904 tons), Ethiopia (190,000 tons), South Sudan (182,153 tons), Uganda (146,000 tons), Mozambique (126,000 tons), Niger (85,062 tons) and Cameroon occupied the 11th position (70,000 tons)
[16]
.
Seeds are used for the nutritional, medicinal, and industrial purposes in Middle East Asia and in Africa
[17]
. The high oil-contain of seeds (about 50%) is the main reason for the cultivation for food (for humans and livestock), pharmaceutical and chemical industries
[17-19]
. In America, Europe, India and Africa, seeds are traditionally used as folk remedy for different disorders such as bowel obstruction, asthma, allergy, and, eye disorders due to its anti-inflammatory, antioxidant and anti-bacterial activities
[17, 20-22]
. In Chinese medicine, sesame seeds are one of the reputed folk medicine used for cure of most symptoms of aging
[17, 20]
. In Algeria, the sesame oil supplementation is recommended to conventional frying oil and to commercial margarine, as alternative source of fatty acids, contributing to the diversification of combined oils
[23]
. Sesame seeds are important grain legume containing high levels of protein, fibres, energy, micronutrients including vitamins B and minerals like copper, iron, calcium, manganese, magnesium, sodium and macro-nutrients whose deficiencies are prevalent in Sub-Saharan African countries and then sesame seeds present a high nutritional value that make it very popular in the diet
[7, 13, 14, 17, 18, 20, 22-24]
. According to the same authors, sesame seeds are rich in lignan-like active ingredients, antioxidant, suitable for cholesterol reduction, blood lipid regulation, liver and kidney protection, cardiovascular system protection, anti-inflammatory, anti-tumor, and other effects, suitable for the human health and the livestock’s nutrition. As an important medicinal and edible homologous food, sesame is used in various aspects of daily life such as food, feed, and cosmetics. The health food applications of sesame are increasing. Sene et al.
[24]
showed that, in eight sesame varieties, protein’s contents ranged from 22.6% to 29.4% whereas that of fats varied from 48.7 to 52.5%, sesame varieties were rich in minerals, calcium being the most representative of all, followed by phosphorus, magnesium, iron, and zinc. The importance of the sesame crop lies in its edible leaves and the seed which is rich in oil (on average 50%), vitamins, proteins (25%), carbohydrates (15%), and minerals. The minerals include calcium, iron, and phosphorus while its vitamin constituents include thiamin, riboflavin and niacin
[25]
. High oil contents of sesame seeds (35 to 60%) were also reported by El Khier et al.
[26]
, Alyemeni et al.
[27]
, Borchani et al.
[28]
and Jimoh et al.
[29]
. Sesame seeds are also a source of essential and sulfur-containing amino acids
[30]
. They are rich in essential fatty acids from the C18 group (linoleic and linolenic acids)
[31]
. In addition, sesame seeds contain many mineral elements and vitamins
[32, 33]
. The strong antioxidant potentials of sesame seeds was highlighted by Dar and Arumugam
[34]
with lignans (sesamolin and sesamin). Sesame seeds are also known to be a source of essential and sulfur-containing amino acids
[35]
. Given its composition of oil, mineral elements, proteins, and antioxidants, sesame (Sesamum indicum L.) is sometimes considered the “queen of oilseeds” and could be used as a food supplement against malnutrition
[36, 37]
.
During the flowering period, flowers of V. unguiculata produce nectar and pollen and very often release in nature a scent attractive to useful and/or harmful animals including insects
[38]
. Harmfull insects on flowers are mostly phytophagous who nibble petals (Hymenoptera, Lepidoptera larvae, Odonata and Orthoptera) and potential pests are sap-sucking insects (Hemiptera and Homoptera) as the case reported in sesame, cowpea, potato and eggplants fields in Cameroon
[39-53]
. Useful insects are predators of harmful ones (natural enemies Coleoptera, Hymenoptera, Neuroptera and Dictyoptera), true pollinators (Hymenoptera), other pollinators (Diptera, Coleoptera and Lepidoptera)
[44]
. For example foragers of the useful Hymenoptera bees (case of Ap. mellifera and Xy. olivacea) face the anthers of the flower, scrape the pollen grains with the metathoracic legs, harvest and carry them in the metathoracic leg baskets (pollen collection). In Cameroon, Ap. mellifera is reported the most frequent floricultural insect on V. unguiculata blooming flowers
[45-53]
. In Benin, the most frequent insect on cowpea flowers was reported as Xy. olivacea
 [54] 
while in Nigeria, Ap. mellifera and Xy. olivacea predominated on V. unguiculata flowers
[55]
. In Ghana, Ap. mellifera and Halictus sp. predominated on cowpea flowers
[56]
. For the nectar collection, bee foragers spread their wings, introduce entirely their head and the proboscis into the flower base to reach the sweet liquid exudates found deep inside the flower
[45]
. When entering the flower, foragers come into contact with the anthers; receive inadvertently pollen grains which adhere to their body, and accidentally release them at the bottom of the corola during the collection of the produced sweet liquid
[45]
. In the world, similar behavior of the bee foragers was reported in Lepidoptera adults and other Hymenoptera Apidae on flowers of many flowering market garden plant species in India including Indonesia
[5-59]
, Pakistan
[60]
, Afrian countries including Egypt
[61]
, Nigeria
[62]
and Cameroon
[39, 45, 46-53, 62-67]
. During the nectar and/or pollen collection, released pollens that escape collection land on the stigma of the flower, facilitating the geitogamy and/or the xenogamy
[68]
.
In Central African countries the sesame production is low compared to the situation in other African Regions and in developing countries and the overall production is insufficient to meet the ever-increasing demand in the cities. Causes of low productions are not fully known but available information points out the influence of abiotic and biotic stresses and socio-economic constraints including the sex and education level of farmers, the lack of improved varieties, insufficient use of fertilizers and low soil fertility, inexperience of farmers, poor access to extension, poor access to credit services, harvesting time, soil conservation, nature of access to land, farmland shortage, access to market, access to irrigation schemes, inadequate phytosanitary control including disease and insect pests, drought, unsuitability of agricultural policies, low soil fertility, the use of infested planting material, high disease and pest infection rates, losses during storage including losses in quality, inappropriate agronomic practices and storage pests. Among the biotitic stresses the useful effect of several animal organisms (bacteria and predators that can protect plants against pests) is counterbalanced by pests (borers, phytophagous and sap-sucking insects). In Cameroon, the sesame production is limited by several factors among which the shortage of agricultural land, the low soil fertility, the poor management of pollinating insects, the pressure from insect pests in the fields and the post-harvest looses in warehouses, are frequently reported
[42-44]
. In natural environments as well as in agro-ecosystems, floricultural insects in general and Apoïdae (Hymenoptera) including Apidae Ap. mellifera, Xy. olivacea and Amegilla spp. have a great ecological indirect impact on the yield production
 [39-52, 59-64]
. The lack of yielding amendment and high quality of seed and absence of resistance to pests and diseases, are known as major problems for the vegetables cultivation.
Although the relationships between flowering plant species and their pollinators have been intensively studied in Cameroon
[17, 42-44]
, no published data exist on the diversity of flower-visiting insects in Bilone. Nevertheless, the control of pest insects as well as useful insects is one of the major constraints to be overcome in sesame cultivation. In the rural areas of Cameroon in general and Bilone in particular, market gardening activities are on the rise, but sesame cultivation is little known by small farmers and it remains practiced by agricultural firms and research centres. Farmers are young, little educated, unassisted and each having a fairly low income. Moreover there is no information concerning the community composition and structure of the flower visiting insects on sesame plants in Bilone (Obala-Cameroon). The purpose of this study is to identify active insects on flowers of the sesame plants, able to reinforce the pollination and influence the quality and/or quantity of agricultural yields.
2. Material and Methods
2.1. Study Site
The study was conducted from May to June 2022 and 2023 at the Bilone agroecological farm (4°10'19.48"N, 11°30'06.53"E; 554 m a.s.l.). Bilone village is located in the northwest of Obala city (Centre Region, Lekie Department) (Figure 1A, 1B and 1C), not far from the N4 national road (Figure 1D). The Obala locality extends between 3°57'0''N, 11°21'0''E and 4°14'0''N, 11°38'0''E, in the forest-savannah ecotone (contact between the Sudanese savannah and the semi-deciduous dense forest)
[69, 70]
. It belongs to the agroecological zone of dense tropical rainforest
[49, 69, 70]
. The plant cover is a mosaic of fallows, home gardens, and cocoa plantations of varying sizes and ages
[50]
. The prevailing climate in Obala and the neighboring areas is a Guinean equatorial savannah
[69]
with dry winter (type Aw) according to the Köppen-Geiger classification with four seasons
[71]
: a short rainy season (mid-March to mid-July of the same year), a short dry season (mid-July to mid-August), a long rainy season (mid-August to mid-November) and a long dry season (mid-November to mid-March of the following year)
[69]
. The rainfall in the Lekie department (around 1,600 mm per year) presents a maximum rainfall in September
[69]
. In Obala and the neighboring areas, the climate parameters experience strong variability both annually and monthly
[69]
. According to the same source of information; the wet season is warm and overcast, the dry season is hot and mostly cloudy, and it is oppressive year round. Over the course of the year, the temperature varies from 20°C in the rainy season to 35°C in the dry season
[69]
. The hot season (from mid-January to mid-April), presents a high average daily temperature (≥31°C) and the hottest month of the year is March (maximum: 31°C; minimum: 22°C)
[69]
.
Figure 1. Localization map of the study site. A: Centre Region in Cameroon
[44]
; B: Lekie department in the Centre Region
[44]
; C: Obala in the lekie department
[44]
; D: Distance from Obala to the Bilone agroecological farm; E: study site at Bilone agroecological farm (Google Earth Pro for windows version 7.3.4.8642).
The cool season (mid-June to September) presents a high average daily temperature below 27°C and the coldest month is July (minimum average: 21°C; maximum: 27°C)
[69]
. According yo the same source of information, the wetter season lasts 8.0 months (mid-March to mid-November) and the month with most wet days is October, an average of 25.2 days presenting at least 1 mm of precipitation. In the dry season (mid-November to mid-March of the following year), the month with the fewest wet days is January. During 2021 and 2022, the temperature generally ranges from 20°C to 32°C and is rarely below 17°C or above 34°. The climate do not show an abnormal variation
[69]
. Soils are ferralitic, thick, homogeneous in appearance and formed on altered original material on which uneven vegetation develops
[69]
. The vegetation is mutilated by humans, notably due to the urban and agricultural development and agricultural operations
[69]
. The major industrial crops in Obala zone include Elaeis guineensis Jacq., 1763 (Arecales: Arecaceae), Musa x paradisiacal L., 1753 (Zingiberales: Musaceae), Threobroma cacao L., 1753 (Malvales: Sterculiaceae) and Coffea arabica L., 1753 (Rubiales: Rubiaceae).
2.2. Experimental Device and Procedure
The investigations were carried out during two years (2022 and 2023) within the campus of the Obala Higher Institute of Agriculture and Management (OHIAM). The experimental plots were created in 1,600 m² area. In the station, several vegetable monoculture plots were created in the vicinity of the sesame plots. Among these neighboring plots were plots of Abelmoschus esculentus (L.) Moench, 1794 (Malvales: Malvaceae), Arachis hypogaea L., 1753 (Fabales: Fabaceae), Capsium annuum L., 1753 (Solanales: Solanaceae), Citrullus lanatus (Thunb.) Matsum. & Nakai, 1916 (Cucurbitales: Cucurbitaceae), Glycine max (L.) Merr., 1917 (Fabales: Fabaceae), Oryza spp. L., 1753 (Poales: Poaceae), Phaseolus vulgaris L., 1753 (Fabales: Fabaceae), Solanum lycopersicum L., 1753 (Solanales: Solanaceae), Solanum tuberosum L., 1753 (Solanales: Solanaceae), Thebroma cacao L., 1753 (Malvales: Sterculiaceae), Solanum sp. (Solanales: Solanaceae), and Zea mays L., 1753 (Cyperales: Poaceae). Seasonal mixed food crop plots were composed with Colocasia esculenta (L.) Schott, 1832 (Arales: Araceae), Mangifera indica L., 1753 (Sapindales: Anacardiaceae), Manihot esculenta Crantz, 1766 (Malpighiales: Euphorbiaceae), Persea americana Mill., 1768 (Laurales: Lauracerae), Psidium guajava L., 1753 (Myrtales: Myrtaceae), Gymnanthemum amygdalinum (Delile) Sch.Bip. ex Walp., 1843 (=Vernonia amydalina Delile) (Asterales: Asteraceae), Xanthosoma sagittifolium (L.) Schott, 1832 (Alismatales: Araceae), and Zea mays L., 1753 (Cyperales: Poaceae). The main wild plants were Bidens pilosa L., 1753 (Asterales: Asteraceae), Lantana camara L., 1753 (Lamiales: Verbenaceae), Mimosa invisa Mart. ex Colla, 1834 (Fabales: Mimosaceae), and Tithonia diversifolia (Hemsl.) A. Gray, 1883 (Asterales: Asteraceae).
Packets of Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) seeds (White and Smooth variety) were obtained from the Institute of Agricultural Research for Development (IRAD/ARID, Nkolbisson station). After the first rains (mid-March of each year), the experimental plots were cleared; ploughed and 15 plots (6 x 5.5 m each) were formed. Subplots were separated from each other by a two-meter wide path and from neighbouring fallows by a two-meter wide safety space. In each subplot, sowing was done in rows (five rows per subplot) and seeds were sown in pockets (10 to 14 seeds per pocket), the spacing being 100 cm on the lines and between the lines. Two weeks after emergence, weeding was done and two plants (the most vigorous) were kept per pocket. From emergence (occurring at the end of March) to the opening of the first flowers (mid-May each year), weeding operations were carried out regularly with a hoe, twice every two weeks. From the start of the flowering period (mid-May each year) to fruit maturity (end of June each year), manual weeding was regularly carried out. Six bee colonies housed in hives with upper bars were positioned between 20 and 24 m from the experimental plots and other colonies were non-inventoried in the vicinity of the study station. During the flowering period, two Sesamum plants were randomly selected each day in each subplot and blooming flowers were checked from 1st blooming day to 13th day (30 flowers a day).
2.3. Capture and Identification of Insects
Throughout the investigation period, 5,241 flowers were monitored in 26 days (13 days in 2022 and 2023 respectively) i.e. in each year, two weeks of the reproductive phase of Se. indicum plants (the week of the early blooming stage and the first week of the mid bloom stage). Then in 2022, a total of 2,883 flowers (55.0% of the total monitored flowers) were monitored (56, 125, 268, 356, 389, 450, 426, 352, 216, 110, 85, 35 and 15 flowers during the 1st to the 13th day respectively). In 2023, a total of 2,358 flowers (45.0%) were monitored (6, 95, 168, 256, 276, 369, 402, 389, 210, 96, 54, 32 and 5 flowers during the 1st to 13th day respectively). Collection sessions were conducted from 22 May to 15 June of each year. Consecutive session days were separated by two days interval. In each day and each year, the blooming flowers were checked during four time periods (9 to 10 a.m., 11 a.m. to 12 p.m., 1 to 2 p.m. and 3 to 4 p.m.). Insects found on the blooming flowers were captured. Products collected by each insect species were determined. Captures were done with bare hands (case of large non-flying insects), using a pair of soft tweezers or a mouth aspirator for entomologists (case of non-flying small insects) or with an entomological net (case of flying insects), following the procedure described by Tchuenguem Fohouo
[42]
. Specimens were preserved in glass pill boxes containing each 70% ethanol, except for adults of Lepidoptera and Odonata which were stored dry. The date and time of the captures were noted.
2.4. Identification of Insect Specimens
Specimens were identified to the family level using keys of Delvare and Aberlenc
[72]
, and Borror and White
[73]
. Bees were identified, to the genera level using the key proposed by Eardley et al.
[74]
, Lecoq
[75]
, Brailovsky
[76]
, Tronquet
[77]
, Taylor
[78]
, and Zettler et al.
[79]
. In order to consider recent developments in the taxonomy, we consulted recent checklists, illustrated catalogues and websites for Diptera
[80-83]
, Hymenoptera
[78, 83-93]
, Lepidoptera
[94-97]
, Neuroptera
[98-100]
, and Orthoptera
[101]
. Identifications were done in the Laboratory of Applied Zoology, Department of Biological Sciences, Faculty of Science, Ngaoundere University where voucher specimens were deposited.
2.5. Data Analysis
Data matrixes of abundance counts of species were constructed in each cultivation campaign and saved using an excel spreadsheet version 2016. Percentages were calculated from the overall total number of specimens or the overall recorded taxa when relevant. Series of abundance counts were presented in terms of mean ± standard error (se) and percentages. Two mean values were compared using the Student t-test from SigmaStat software 2.03 (SPSS, Inc., Chicago, IL), when relevant and when normality and equal variance tests passed. Otherwise the non-parametric Wilcoxon test (paired series) or Mann-Whitney test (independent series) was used. Comparison of two frequencies was done using the Fisher’s exact-test from StatXact software 3.1. The link between the occurrence of insects and climatic parameters (temperature and air humidity) was evaluated by determining the Pearson correlation. Regression equations were set up when relevant and tested using ANOVA procedure.
Alpha diversity analysis allowed the determination of indices using PAST 3.05 software
[102]
: the absolute abundance of the ith species ni, the sample size n (sum of ni), the maximum abundance nmax, the relative abundance of the ith species fi=ni/n, the observed species richness S (total number of the collected species), the Shannon-Weaver index H’, the maximum Shannon-Weaver index H’max=ln(S), the Simpson’s index D (D=0 for high diversity), the Margalef’s index Mg=(S-1)/ln(n) with 0≤Mg≤+∞ (Mg=0 for a low species richness). The ‘true’ theoretical richness T was determined using six non-parametric estimators from EstimateS software Version 9.1.0
[103]
: the Abundance Coverage-based Estimator (ACE), Chao 1, Chao 2, the Incidence Coverage-based Estimator (ICE), Jackniffe estimator of order 1 (Jack 1), and Boostrap Mean. For each estimator, the sampling success was determined as SE=(S/T)*100. Comparison of the species richness was performed using the individual rarefaction procedure and pair wise comparison of diversities (H’ and D) was performed using the Student t-test from PAST 3.05 software
[102]
. The Pielou’s evenness index J=H’/H’max and the Hill’s diversity numbers (N1=eH’ and N2=1/D) were determined. The richness ratio d=S/n with 0≤d≤1, confirmed the quality of the species richness (d close to null for low species richness and d close to one for high species richness). The degree of dominance by a few species was evaluated using the Berger-Parker index IBP=nmax/n with 0≤IBP≤1 (IBP close to null for equally abundances).
For the beta diversity, the overall species covariance was evaluated using the Schluter’s procedure
[104]
and between species correlations was determined using the Kendall’s tau coefficient. The dissimilarity between the two years was evaluated using the Bray-Cutis index
[105]
. The rank abundance plotting illustrated the shape of the species abundance distributions (SADs). Species were ranged in decreasing order of abundance and the absolute value of the Bravais-Pearson correlation between ranks i and Log2(ni) maked it possible to assess the adjustment of Motomura's law to observed data. We tested five commonly used theoretical models
[106]
to fit the curves, using the package vegan of R 3.4.1 software: Broken-stick (BS), log-linear (LL), Log-normal (LN), Zipf (Z) and Zipf-Mandelbrot (ZM). The best fitted model presented the lowest value of the Akaike Information Criteria (AIC) or the lowest Bayesian Information Criteria (BIC)
[107]
. For each selected theoretical model, the estimated sample size n* was adjusted to the observed sample size n using the correction factor c=n/n*, and the corrected model was given. BS model has a single parameter x (average abundance). LL or GM model corresponds to the linear regression Log2(ni)=a(i)+b or ni=c*2b*(2a)i where i represents the rank of the species in decreasing order of abundance, ni is the abundance of the ith species, a and b represent the slope and the elevation of the regression respectively. LL model depends on the maximum abundance of the top-ranking species n1 and the Motomura environmental constant m (antilogarithm of the regression slope a, 0≤m≤1) representing the rate of decrease in abundance by rank. The LN model corresponds to the linear regression Log2(ni)=a(Pi)+b or ni=c*2b*(2a)Pi where Pi represents the probit of the ith species. For a species of rank i, the cumulative percentage linked to the rank was determined as ki=100(i+0.5)/(S+1) when S was odd or ki=100((i+1)+0.5)/(S+1) when S was even. Probit were determined using the package “Ecotoxicology” from R 3.4.1 software. Parameters of LN were the maximum abundance n1, the mean of the lognormal distribution x, the standard deviation of the lognormal distribution σ and the Preston’s environmental constant (rate of decrease in abundance by rank) m’=square root of 1/σ. Z model is based on the Zipf’s law based on two statistics
[108]
: Q as the scaling parameter (normalizing constant), and γ (gamma) as the average probability of the appearance of a species
[108]
. Zipf's law
[109]
is frequently applied in animal and plant ecology to characterize SADs. ZM is a generalized model in which a new parameter β (beta) is added. Marquardt’s nonlinear least squares algorithm
[110, 111]
was used when relevant to estimate β, γ and 1/γ parameters (fractal dimension of the distribution of individuals among species).
3. Results
3.1. Inventory and Abundances of Insects
A total of 1,703 adult insects collected in 2022 and 2023, belonged to five orders, 12 families (1,703 specimens, 8 to 721 specimens, mean±se: 142±71 specimens, Me=22 specimens), 18 genera and 19 species (Table 1). Orders were Diptera Linnaeus, 1758, Hymenoptera Linnaeus, 1758, Lepidoptera Linnaeus, 1758, Neuroptera Linnaeus, 1758, and Orthoptera Latreille, 1793 (Table 1). Hymenoptera was the most family-rich order (six families) followed by Diptera and Lepidoptera (two families each). Neuroptera and Orthoptera were rare (one Family each) (Table 1). Calliphoridae, Eumenidae and Halictidae were not recorded in 2022. In each year, Hymenoptera was mostly recorded (91.5%). Other orders were rare (Table 1). Apidae Latreille, 1802 was mostly recorded (42.3% of the collection) followed by Formicidae Latreille, 1809 (34.1%), Megachilidae Latreille, 1802 (11.6%). Other families were rare (Table 1). Mean or median occurrences in 2022 (five orders, 10 families, 677 specimens, 4-290 specimens, mean abundance ± se: 68±33 specimens, median: Me=11 specimens) was not different from the records in 2023 (five orders, 11 families, 1,026 specimens, three to 431 specimens, 93±46 specimens, Me=15 specimens) (Student t-test: t=-0.443, df=19, p=0.663; Mann-Whitney test: T=102.500, p=0.622). Calliphoridae (Diptera) and Ascalapidae (Neuroptera) were highly abundant in 2023 campaign than the 2022 one while it was the contrary in Eumenidae, Halictidae, Megachilidae, the pooled data of Hymenoptera and the overall pooled data (Table 1). Between the two years, the difference was not significant in Acrididae, Apidae, pooled Diptera, Formicidae, pooled Lepidoptera, Muscidae, Nymphalidae, Pieridae and Vespidae (Table 1). The most species-rich family was Formicidae (four species), followed by Apididae (three species), Megachilidae and Nymphalidae were represented each by two species. Acrididae, Ascalapidae, Calliphoridae, Eumenidae, Halictidae, Muscidae, Pieridae, and Vespidae were represented each by one species (Table 2). Calliphora vicina Robineau-Desvoidy, 1830 (Diptera: Calliphoridae) was recorded exclusively in 2022 (Table 2). Two useful species: the exotic Eumenidae Delta sp. and the afrotropical predator Ascalaphus africanus (Ascalapidae) were recorded as well as the phytophagous Acrididae Pe. carnapi. Potential pests (Nymphalidae, Pieridae, and Acrididae) cumulatively represented 3.1%.
Table 1. Absolute and relative abundance of insect orders and families collected on flowers of Sesamum indicum L. (1753) (Pedaliaceae).

Orders / Families

Campaign

2022 (%)

2023 (%)

Total (%)

2022 vs. 2023: Fisher’s exact test

Diptera Linnaeus, 1758

Calliphoridae Hough (d), 1899

8 (0.5)

-

8 (0.5)

χ²=8.607; df=1; p=0.008 *

Muscidae Latreille, 1802

29 (1.7)

39 (2.3)

68 (4.0)

χ²=0.273; df=1; p=0.615 ns

Total

37 (2.2)

39 (2.3)

76 (4.5)

χ²=2.645; df=1; p=0.119 ns

Hymenoptera Linnaeus, 1758

Apidae Latreille, 1802

290 (17.0)

431 (25.3)

721 (42.3)

χ²=0.116; df=1; p=0.764 ns

Eumenidae Leach, 1815

-

12 (0.7)

12 (0.7)

χ²=9.326; df=1; p=0.003 *

Formicidae Latreille, 1809

216 (12.7)

364 (21.4)

580 (34.1)

χ²=2.313; df=1; p=0.130 ns

Halictidae Thomson, 1869

-

37 (2.23)

37 (2.2)

χ²=34.026; df=1; p=9.6x10-9 *

Megachilidae Latreille, 1802

97 (5.7)

101 (5.9)

198 (11.6)

χ²=7.878; df=1; p=0.005 *

Vespidae Latreille, 1802

4 (0.2)

7 (0.4)

11 (0.6)

χ²=0.070; df=1; p=1.00 ns

Total

607 (35.6)

952 (55.9)

1,559 (91.5)

χ²=6.101; df=1; p=0.026 *

Lepidoptera Linnaeus, 1758

Nymphalidae Rafinesque, 1815

9 (0.5)

13 (0.8)

22 (1.3)

χ²=0.045; df=1; p=1.00 ns

Pieridae Swainson, 1820

5 (0.3)

3 (0.2)

8 (0.5)

χ²=1.734; df=1; p=0.278 ns

Total

14 (0.8)

16 (0.9)

30 (1.8)

χ²=0.654; df=1; p=0.456 ns

Neuroptera Linnaeus, 1758

Ascalapidae Rambur, 1842

12 (0.7)

4 (0.2)

16 (0.9)

χ²=8.075; df=1; p=0.008 *

Orthoptera Latreille, 1793

Acrididae MacLeay, 1821

7 (0.4)

15 (0.9)

22 (1.3)

χ²=0.532; df=1; p=0.516 ns

Global

677 (39.8)

1,026 (60.2)

1,703 (100.0)

χ²=142.98; df=1; p=5.3x10-33 *
ns: not significant difference (p>0.05); *: significant difference (p<0.05)
3.2. Alpha Diversity of the Insects’ Assemblages
The numbers of species recorded in 2022 and 2023 were close to each other and revealed in each case, low species richness (richness ratio close to 0) (Table 3A). The species richness was low in 2022 (16 species; Margalef index: Mg=2.301; richness ratio: d=0.024) and high in 2023 (18 species; Mg=2.452; d=0.018) and in the pooled years (19 species; Mg=2.419; d=0.011) (Table 3A). The sampling success was maximal (100.0%), suggesting no rare species escaped (Table 3B). In each year, a high diversity of the assemblage was noted (Shannon index close to the maximum; Table 3C). A low dominance by a few species was noted (Berger-Parker index inferior to the median value; Table 3D).
Based on the Hill's N1 and N2 indexes, the number of simply abundant species were close to the number of co-dominants and values of the Hill's ratio were very close to one (Table 3D), corroborating a low dominance of the assemblages by a few insect species. The number of rare species was 7 species in 2022, height species in 2023 and nine species in the pooled years (Table 3D). A high even assemblage was noted (Pielou’s index close to one; Table 3E). The variation in the diversity indexes was not significant between the two years. The rank-abundance plotting presented in the pooled campaigns, a concave appearance suggesting the presence of co-dominants (Figure 2A). The similar shape was noted in the species distribution of abundances (SADs) recorded in 2022 and 2023 (Figure 2B and 2C). The individual rarefaction curves plotted for the two campaigns and the pooled campaigns approached species saturation plateaus with similar slopes (Figure 2D). The curve observed in 2022 was situated below records in 2023 and the pooled years, suggesting lowest species richness in 2022. A high species richness was noted in 2023 and in the pooled years (Figure 2D). For a standard sample of 661 specimens, the settlement in the pooled years was most diversed (E(Sn=661)=19±0 species), followed by 2023 (E(Sn=661)=18±0 species), and lastly the records in 2022 (E(Sn=661)=16±0 species).
Based on the Hill’s first order diversity number N1 (see Table 3) and the rank-abundance plotting (Figure 2), the number of simply abundant species varied from 9 species (47.4% of the total species richness) in 2022 to 10 species (52.6%) in 2023 and the pooled years respectively. Camponotus maculatus was simply abundant in 2023 and in the pooled years. Nine species were simply abundant in each year and in the pooled years. These species were Am. calens, Ap. mellifera, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Ph. megacephala, and Xy. olivacea. Based on the Hill’s second order diversity number N2 (Table 3) and the rank-abundance plotting (Figure 2), seven species were co-dominants (36.8% of the total species richness) in all cases. Two species (Am. calens and Me. cincta) were co-dominants in 2022 and in the pooled campaign. Two species (Ca. maculatus and Me. kamerunensis) were co-dominants exclusively in 2023. Five species (Ap. mellifera adansonii, Pa. longicornis, Ph. Megacephala, My. opaciventris and Xy. olivacea) were co-dominants in each year and in the pooled years. Camponotus maculatus was rare exclusively in 2022. Calliphora vicina was rare in 2022 and in the pooled years. Three species (Bi. dorothea, Delta sp. and La. hancocki) were rare in 2023 and in the pooled years. Five species (Ac. acerata, As. africanus, Ct. florella, Pe. carnapi and Sy. conuta) were rare in both years and in the pooled years.
3.3. Abundance Distributions (SADs)
Adjustment of the SADs to the five commonly known theoretical models showed that the fit was of excellent quality in 2022 (r=-0.990, p=3.6x10-13, 16 species), of satisfactory quality in 2023 (r=-0.977, p=4.1x10-12, 18 species), and of excellent quality in the pooled campaigns (r=-0.986, p=8.9x10-15, 19 species). On the base of the AIC and BIC values (Table 4) and the SAD plotting (Figure 2A, 2B and 2C), the log-linear model (LL) best fitted the insect assemblage in the 2022 with a high Motomura’s environmental constant close to one (maximum abundance: n1=194 specimens; sample size: n=677 specimens; species richness: S=16 species; log-linear regression slope: a=(-0.110±0.004; Student test t=-25.638; p<0.001); Motomura’s environmental constant: m=0.777; elevation of the regression: b=(2.278±0.041; Student test t=55.046; p<0.001); ANOVA log-linear regression: F(1, 14)=657.291, p<0.001; deviance: 23.191; correction factor: 1.045; corrected LL model: ni=198.031*(0.777)i with i as the rank of species, arranged in descending order of abundance. The settlement in 2023 best fitted the lognormal model (LN) with a high value of the Preston’s environmental constant (deviance: 38.368; n1=327; mean of the lognormal distribution: x=4.80; standard deviation of the lognormal distribution: σ=1.896; slope of Log2(ni)=f(Pi): a=1.207; elevation: b=-0.808; Preston’s environmental constant: m’=0.726; correction factor: 8.0x10-5; corrected model: ni=0.810(2.31)Pi with Pi as the probit of the ith species.
Table 2. Absolute and relative abundance of the insect species collected on flowers of Sesamum indicum L. (1753) (Pedaliaceae).

Order/Family

Species name

Product

Origin, references

Campaign

2022 (%)

2023 (%)

Pooled (%)

Diptera

Calliphoridae

Calliphora vicina Robineau-Desvoidy, 1830

Nectar

NA, MS, a

8 (0.5)

-

8 (0.5)

Muscidae

Musca domestica Linnaeus, 1758

Nectar

ME, MS, b, c

29 (1.7)

39 (2.3)

68 (4.0)

Hymenoptera

Apidae

Amegilla calens (Lepeletier De Saint-Fargeau, 1841)

Nectar, Pollen

AF, d, e

59 (3.5)

46 (2.7)

105 (6.2)

Apis mellifera adansonii Latreille, 1804

Nectar

AF, e

194 (11.4)

327 (19.2)

521 (30.6)

Xylocopa olivacea (Fabricius 1778)

Nectar, Pollen

AF, f

37 (2.2)

58 (3.4)

95 (5.6)

Eumenidae

Delta sp. Saussure, 1855

Predator

OW, Useful, g, k

-

12 (0.7)

12 (0.7)

Formicidae

Camponotus maculatus (Fabricius, 1782)

Nectar

AF, h, i

6 (0.4)

53 (3.1)

59 (3.5)

Myrmicaria opaciventris Emery, 1893

Nectar

AF, h, i

80 (4.7)

71 (4.2)

151 (8.9)

Paratrechina longicornis (Latreille, 1802)

Nectar

AF, h, i

90 (5.3)

120 (7.0)

210 (12.3)

Pheidole megacephala (Fabricius, 1793)

Nectar

AF, h, i, j

40 (2.3)

120 (7.0)

160 (9.4)

Halictidae

Lasioglossum hancocki (Cockerell 1945)

Nectar, Pollen

AF, m

-

37 (2.2)

37 (2.2)

Megachilidae

Megachile cincta (Fabricius, 1781)

Nectar

AF, l

81 (4.8)

38 (2.2)

119 (7.0)

Me. kamerunensis Friese, 1922

Nectar, Pollen

AF, l

16 (0.9)

63 (3.7)

79 (4.6)

Vespidae

Synagris conuta (Linnaeus, 1758)

Nectar, Pollen

AF, n, o

4 (0.2)

7 (0.04)

11 (0.6)

Lepidoptera

Nymphalidae

Acraea acerata Hewitson, 1874

Nectar

AF, Plant pest, p

9 (0.5)

6 (0.4)

15 (0.9)

Bicyclus dorothea (Cramer, 1779)

Nectar

AF, p, q, r

-

7 (0.4)

7 (0.4)

Pieridae

Catopsilia florella (Fabricius, 1775)

Nectar

AF, p, s

5 (0.3)

3 (0.2)

8 (0.5)

Neuroptera

Ascalapidae

Ascalaphus africanus (McLachlan, 1871)

Predator

AF, Useful, t, u, v

12 (0.7)

4 (0.2)

16 (0.9)

Orthoptera

Acrididae

Pteropera carnapi Ramme, 1929

Phytophagous

AF, Plant pest, w

7 (0.4)

15 (0.9)

22 (1.3)

Total

677 (39.8)

1,026 (60.2)

1,703(100.0)
AF: Afrotropical origin; MS: Myiasigenic species; ME: Middle East (Asia); NA: North America origin; OR: oriental origin; OW: Old World; a:
[81]
; b:
[80]
; c:
[82]
; d:
[90]
; e:
[89]
; f:
[95]
; g:
[84]
; h:
[78]
; i:
[83]
; j:
[86]
; k:
[91]
; l:
[87]
; m:
[88]
; n:
[93]
; o:
[85]
; p:
[92]
; q:
[94]
; r:
[96]
; s:
[97]
; t:
[99]
; u:
[98]
; v:
[100]
; w:
[101]
.
Table 3. Alpha diversity indices of the floricultural insects on flowers of Sesamum indicum L. (1753) (Pedaliaceae).

Statistical indices

Campaign

I. 2022

II. 2023

III. Pooled years

A. Species richness indices

Sample size n (%)

677 (39.8)

1,026 (60.2)

1,703 (100.0)

Observed species richness S

16

18

19

Maximum abundance nmax

194

327

521

Margalef’s index Mg

2.301

2.452

2.419

Richness ratio d=S/n

0.024

0.018

0.011

B. Non-parametric estimation of the "true" species richness

ACE (SE=100*S/ACE)

16 (100.0)

18 (100.0)

19 (100.0)

ICE (SE=100*S/ICE)

16 (100.0)

18 (100.0)

19 (100.0)

Chao1 (SE=100*S/Chao1)

16 (100.0)

18 (100.0)

19 (100.0)

Chao2 (SE=(100*S/Chao2)

16 (100.0)

18 (100.0)

19 (100.0)

Jack.1

16 (100.0)

18 (100.0)

19 (100.0)

Boostrap Mean

16 (100.0)

18 (100.0)

19 (100.0)

C. Species diversity indices

Shannon-Weaver H’

2.232

2.294

2.329

Maximum Shannon-Weaver H’max=ln(S)

2.773

2.890

2.944

Simpson index D

0.145

0.150

0.143

D. Species dominance indices

Berger-Parker dominance index IBP=nmax/n

0.287

0.319

0.306

Hill’s first order diversity number N1=eH’

9.318

9.914

10.268

Hill’s second order diversity number N2=1/D

6.878

6.667

6.983

Hill’s ratio: Hill=N2/N1

0.738

0.672

0.680

Estimated observed rare species: Chao1-N1

7

8

9

E. Evenness index

Pielou’s index J=H’/H’max

0.805

0.794

0.791

Comparison of the species diversity indices: I vs. II (Student t-test):

Shannon-Weaver index H’: t=-1.332; df=1,532.1; p=0.183 ns;

Simpson index D: t=-0.433; df=1,635.1; p=0.665 ns;
ns: not significant difference (p>0.05); SE: sampling effort; ACE: Abundance Coverage-based Estimator; ICE: Incidence Coverage-based Estimator; Chao1: first order Chao index; Chao2: second order Chao index; Jack.1: first order Jackniffe estimator.
Figure 2. Rank-frequency diagrams of the collected insects in the pooled campaigns (A), in 2022 (B) and 2023 (C) showing species in order of numerical dominance. The species rarefaction curves (D) (estimated species richness as a function of the sample size variation) showed the low species richness variation in 2022 and the high species richness in the pooled campaigns.
The pooled campaigns fitted the LN model with a high value of the Preston’s environmental constant (deviance: 53.367; n1=521; mean of the lognormal distribution: x=5.43; standard deviation: σ=1.868; slope of Log2(ni)=f(Pi): a=1.152; elevation: b=0.057; Preston environmental constant: m’=0.732; correction factor: 1.5; corrected model: ni=1.5(2.22)Pi with Pi as the probit of the ith species.
3.4. Beta Diversity of the Insects Assemblages
Based on the species composition, although a few cosmopolitan species were sampled, a high level of dissimilarity was noted between 2022 and 2023 campaigns (Bray-Curtis index: BC=0.694), between 2023 and the pooled campaigns (BC=0.752) and it was of median level between 2022 and the pooled campaigns (BC=0.569). Species were recorded on 1,066 flowers (5,241 checked flowers: 20.3%): 294 flowers (5.7%) in 2022 and 772 flowers (14.7%) in 2023. Overall, insects exhibited in 2022, a positive net association in presence/absence data (VR>1) (variance ratio: VR=2.485, statistic: W=7,165.691, df=2,882, p<0.001). It was the same in 2023 (VR=6.513, W=15,357.456, df=2,357, p<0.001) and in the pooled years (VR=8.592, W=45,029.530, df=5,240, p<0.001). A few negatively correlated species (mutual repulsion) and several positively correlated ones (mutual repulsion) were noted. A negative correlation was noted between Calliphoridae Cl. vicina and Apidae Ap. mellifera adansonii (Table 5). The Nymphalidae Ac. acerata was positively correlated with 16 species: Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta (Table 5). The Apidae Am. calens was positively correlated with 16 species (Ac. acerata, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta) (Table 5). Apis mellifera adansonii (Apidae) and Xy. olivacea were positively correlated (Table 5). Ascalaphus africanus (Ascalapidae) was positively correlated with 16 species: Ac. acerata, Am. calens, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapis, Ph. megacephala, and Sy. conuta (Table 5). Bicyclus dorothea (Nymphalidae) was positively correlated with 16 species: Ac. acerata, Am. calens, As. africanus, Ca. maculatus, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, Sy. cornuta, and Xy. olivacea (Table 5).
Table 4. Values of the Akaike Information Criteria and the Bayesian Information Criteria for the adjusted theoretical models of the species abundance distributions.

SAD theoretical model

Deviance; AIC (BIC)

I. 2022 campaign 16 species; 677 specimens

II. 2023 campaign 18 species; 1,026 specimens

III. Pooled years 19 species; 1,703 specimens

McArthur’s Broken-Stick (BS)

49.920; 129.097 (129.097)

115.881; 209.095 (209.095)

191.935; 298.518 (298.52)

Motomura’s Log-linear (LL)

23.191; 104.368 (105.140) *

106.250; 201.464 (202.354)

126.679; 235.261 (236.21)

Preston’s Log-normal (LN)

26.090; 109.267 (110.812)

38.368; 135.582 (137.363) *

53.367; 163.95 (165.84) *

Zipf (Z)

70.894; 154.071 (155.616)

91.644; 188.857 (190.638)

155.56; 266.142 (268.03)

Zipf-Mandelbrot (ZM)

21.740; 106.916 (109.234)

79.141; 178.354 (181.025)

103.945; 216.528 (219.36)
AIC: Akaike Information Criteria; BIC: Bayesian Information Criteria; SAD: Species Abundance Distribution; S: observed species richness; n: sample size; *: the best fitted theoretical model
Camponotus maculatus (Formicidae) was positively correlated with 13 species: Ac. acerata, Am. calens, As. africanus, Bi. dorothea, La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapis, Ph. megacephala, and Sy. cornuta (Table 5). Calliphora vicina (Calliphoridae) was positively correlated with 12 species: Ac. acerata, Am. calens, As. africanus, Ct. florella, Me. cincta, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta (Table 5). Other correlations were not significant (Table 5). Catopsilia florella (Pieridae) was positively correlated with 15 species: Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Cl. vicina, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapis, Ph. megacephala, and Sy. cornuta (Table 5). Delta sp. (Eumenidae) was positively correlated with 16 species: Ac. acerata, Am. calens, As. africanus, Ca. maculatus, Ct. florella, Bi. dorothea, La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, Xy. olivacea, and Sy. conuta (Table 5). Lasioglossum (Ipomalictus) hancocki (Halictidae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Ct. florella, Delta sp., Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, Sy. cornuta, and Xy. olivacea) (Table 5). Megachile (Chalicodoma) cincta (Megachilidae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta) (Table 5). Megachile kamerunensis (Megachilidae) was positively correlated with 15 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta) (Table 5). Musca domestica (Diptera) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, My. opaciventris, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta) (Table 5). Myrmicaria opaciventris (Formicidae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, Pa. longicornis, Pe. carnapi, Ph. megacephala, and Sy. cornuta) (Table 5).
Table 5. Kendall tau τ correlation between 19 species recorded in 1,066 flowers.

Species 1/species 2

tau τ

p-value

Species 1/species 2

tau τ

p-value

Acraea acerata

Apis mellifera adansonii

Ascalaphus africanus

0.962

6x10-12 *

Me. cincta

-0.046

0.739 ns

Bicyclus dorothea

0.622

8x10-6 *

Me. kamerunensis

-0.070

0.617 ns

Catopsilia florella

0.593

2x10-5 *

My. opaciventris

-0.103

0.463 ns

Pteropera carnapi

0.843

2x10-9 *

Pa. longicornis

-0.056

0.689 ns

Amegilla calens

Pe. carnapi

-0.082

0.559 ns

Acraea acerata

0.660

2x10-6 *

Ph. megacephala

-0.070

0.617 ns

Apis mellifera adansonii

-0.135

0.335 ns

Sy. conuta

-0.103

0.462 ns

As. africanus

0.655

3x10-6 *

Xy. olivacea

0.639

5x10-6 *

Bi. dorothea

0.426

0.002 *

As.africanus

Camponotus maculatus

0.648

3x10-6 *

Pe. carnapi

0.786

2x10-8 *

Ct. florella

0.403

0.004 *

Bi. dorothea

Delta sp.

0.449

0.001 *

As. africanus

0.596

2x10-5 *

Lasioglossum (Ipomalictus) hancocki

0.449

0.001 *

Ct. florella

0.449

0.001 *

Megachile (Chalicodoma) cincta

0.565

5x10-5 *

Pe. carnapi

0.758

6x10-8 *

Me. (Chalicodoma) kamerunensis

0.744

1x10-7 *

Camponotus maculatus

Myrmicaria opaciventris

0.958

7x10-12 *

Ac. acerata

0.685

9x10-7 *

Paratrechina longicornis

0.775

3x10-8 *

As. africanus

0.634

6x10-6 *

Pe. carnapi

0.570

5x10-5 *

Camponotus maculatus

Pheidole megacephala

0.759

5x10-8 *

Bi. dorothea

0.634

6x10-6 *

Synagris conuta

0.545

9x10-5 *

Ct. florella

0.206

0.139 ns

Xy. olivacea

-0.015

0.915 ns

La. hancocki

0.663

2x10-6 *

Apis mellifera adansonii

Me. cincta

0.399

0.004 *

Ac. acerata

-0.197

0.158 ns

Me. kamerunensis

0.885

2x10-10 *

As. Africanus

-0.222

0.111 ns

My. opaciventris

0.659

2x10-6 *

Bi. dorothea

0.049

0.723 ns

Pa. longicornis

0.709

4x10-7 *

Ca. maculatus

0.036

0.798 ns

Pe. carnapi

0.827

3x10-9 *

Ct. florella

-0.263

0.059 ns

Ph. megacephala

0.866

6x10-10 *

Delta sp.

0.066

0.637 ns

Sy. conuta

0.465

0.001 *

La. hancocki

0.066

0.637 ns
Table 5. Continued.

Species 1/species 2

tau τ

p-value

Species 1/species 2

tau τ

p-value

Calliphora vicina

La. hancocki

Ac. acerata

0.652

3x10- *

Ac. acerata

0.652

3x10-6 *

Am. calens

0.472

0.001 *

As. africanus

0.596

2x10-5 *

Ap. mellifera adansonii

-0.329

0.018 *

Bi. dorothea

0.959

6x10-12 *

As. africanus

0.715

3x10-7 *

Ct. florella

0.408

0.003 *

Bi. dorothea

-0.082

0.559 ns

Me. cincta

0.449

0.001 *

Ca. maculatus

0.236

0.091 ns

La. hancocki

Ct. florella

0.490

5x10-4 *

Me. kamerunensis

0.573

4x10-5 *

Delta sp.

-0.082

0.559 ns

Pe. carnapi

0.791

1x10-8 *

La. hancocki

-0.082

0.559 ns

Sy. conuta

0.746

9x10-8 *

Me. cincta

0.426

0.002 *

Me. cincta

Me. kamerunensis

0.520

2x10-4 *

Ac. acerata

0.627

7x10-6 *

Musca domestica

0.359

0.010 *

As. africanus

0.622

8x10-6 *

My. opaciventris

0.393

0.005 *

Bi. dorothea

0.426

0.002 *

Pa. longicornis

0.474

0.001 *

Ct. florella

0.403

0.004 *

Ph. megacephala

0.546

9x10-5 *

Me. kamerunensis

0.518

2x10-4 *

Pe. carnapi

0.320

0.022 *

Pe. carnapi

0.532

1x10-4 *

Sy. comuta

0.373

0.008 *

Sy. conuta

0.545

9x10-5 *

Xy. olivacea

-0.265

0.057 ns

Me. kamerunensis

Ct. florella

Ac. acerata

0.822

4x10-9 *

As. africanus

0.670

2x10-6 *

As. africanus

0.797

1x10-8 *

Pe. carnapi

0.286

0.040 *

Bi. dorothea

0.546

9x10-5 *

Delta sp.

Ct. florella

0.493

4x10-4 *

Ac. acerata

0.652

3x10-6 *

Pe. carnapi

0.725

2x10-7 *

As. africanus

0.596

2x10-5 *

Sy. conuta

0.653

3x10-6 *

Bi. dorothea

0.959

6x10-12 *

Mu. domestica

Delta sp.

Ac. acerata

0.648

4x10-6 *

Ca. maculatus

0.663

2x10-6 *

Am. calens

0.758

6x10-8 *

Ct. florella

0.408

0.003 *

Ap. mellifera adansonii

-0.131

0.348 ns

La. hancocki

1.000

8x10-13 *

As. africanus

0.609

1x10-5 *

Me. cincta

0.449

0.001 *

Bi. dorothea

0.487

5x10-4 *

Me. kamerunensis

0.573

4x10-5 *

Ca. maculatus

0.636

5x10-6 *

My. opaciventris

0.462

0.001 *

Ct. florella

0.359

0.010 *

Pa. longicornis

0.439

0.002 *

Delta sp.

0.510

3x10-4 *

Pe. carnapi

0.791

1x10-8 *

La. hancocki

0.510

3x10-4 *

Ph. megacephala

0.573

4x10-5 *

Me. cincta

0.346

0.013 *

Sy. conuta

0.746

9x10-8 *

Me. kamerunensis

0.688

8x10-7 *
Table 5. Continued.

Species 1/species 2

tau τ

p-value

Species 1/species 2

tau τ

p-value

My. opaciventris

0.734

1x10-7 *

Ph. megacephala

Pa. longicornis

0.702

5x10-7 *

Ac. acerata

0.803

9x10-9 *

Pe. carnapi

0.602

2x10-5 *

As. africanus

0.797

1x10-8 *

Ph. megacephala

0.673

1x10-6 *

Bi. dorothea

0.546

9x10-5 *

Sy. conuta

0.529

2x10-4 *

Ct. florella

0.520

2x10-4 *

Xy. olivacea

0.005

0.971 ns

La. hancocki

0.573

4x10-5 *

My. opaciventris

Me. cincta

0.534

1x10-4 *

Ac. acerata

0.604

2x10-5 *

Me. kamerunensis

0.983

2x10-12 *

As. Africanus

0.598

2x10-5 *

Pe. carnapi

0.703

5x10-7 *

Bi. dorothea

0.439

0.002 *

Sy. conuta

0.675

1x10-6 *

Ct. florella

0.393

0.005 *

Sy. conuta

My. opaciventris

Ac. acerata

0.751

8x10-8 *

La. hancocki

0.462

0.001 *

As. africanus

0.804

8x10-9 *

Me. cincta

0.547

9x10-5 *

Bi. dorothea

0.780

2x10-8 *

Me. kamerunensis

0.754

7x10-8 *

Ct. florella

0.814

6x10-9 *

Pa. longicornis

0.784

2x10-8 *

Pe. carnapi

0.573

4x10-5 *

Pe. carnapi

0.524

2x10-4 *

Xy. olivacea

Ph. megacephala

0.769

4x10-8 *

Ac. acerata

0.039

0.783 ns

Sy. conuta

0.537

1x10-4 *

As. africanus

0.013

0.926 ns

Pa. longicornis

Bi. dorothea

0.309

0.027 *

Ac. acerata

0.654

3x10-6 *

Ca. maculatus

0.198

0.156 ns

As. africanus

0.649

3x10-6 *

Ct. florella

-0.009

0.949 ns

Bi. dorothea

0.416

0.003 *

Delta sp.

0.327

0.019 *

Ct. florella

0.404

0.004 *

La. hancocki

0.327

0.019 *

La. hancocki

0.439

0.002 *

Me. cincta

0.205

0.143 ns

Pa. longicornis

Me. kamerunensis

0.098

0.482 ns

Me. cincta

0.534

1x10-4 *

My. opaciventris

0.020

0.886 ns

Me. kamerunensis

0.814

6x10-9 *

Pa. longicornis

0.115

0.410 ns

Ph. megacephala

0.829

3x10-9 *

Pe. carnapi

0.160

0.250 ns

Pe. carnapi

0.562

6x10-5 *

Ph. megacephala

0.098

0.482 ns

Sy. conuta

0.537

1x10-4 *

Xy. olivacea

Sy. conuta

0.140

0.317 ns
ns: not significant correlation (p≥0.05); *: significant correlation (p<0.05). Significant correlations are in bold.
Paratrechina longicornis (Formicidae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pe. carnapi, Ph. megacephala, and Sy. cornuta) (Table 5). Pteropera carnapi (Acrididae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Ph. megacephala, and Sy. cornuta) (Table 5). Pheidole megacephala (Formicidae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, My. opaciventris, Pa. longicornis, Pe. carnapis, and Sy. cornuta) (Table 5). Synagris cornuta (Vespidae) was positively correlated with 16 species (Ac. acerata, Am. calens, As. africanus, Bi. dorothea, Ca. maculatus, Cl. vicina, Ct. florella, Delta sp., La. hancocki, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Pe. carnapi, and Ph. megacephala) (Table 5). Xyloxopa olivacea (Apidae) was positively correlated with four species (Ap. mellifera adansonii, Bi. dorothea, Delta sp., and La. hancocki) (Table 5).
4. Discussion
4.1. Species Richness, Diversity, Abundances
The study carried out on the flower-visiting insects fauna revealed that on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) plant flowers these insects belonged to five orders, 12 families, 18 genera, and 19 species. Hymenoptera represented more than 91.5% of the collected insects. Other orders were rare and represented each by less than 5% of the total collection: Diptera (4.5%), Lepidoptera (1.8%), Neuroptera (0.9%), and Orthoptera (1.3%). Apidae was the most recorded (42.4%) followed by Formicidae (34.1%), Megachilidae (11.6%). Other families were rare and represented each by less than 5%: Acrididae (1.3%), Ascalapidae (0.9%), Calliphoridae (0.5%), Eumenidae (0.7%), Halictidae (2.2%), Muscidae (4.0%), Nymphalidae (1.3%), Pieridae (0.5%), and Vespidae (0.6%). The most recorded species was Apis mellifera adansonii (Apidae) (30.6%), followed very far by Paratrechina longicornis (Formicidae) (12.3%), Pheidole megacephala (Formicidae) (9.4%), Myrmicaria opaciventris (Formicidae) (8.9%), Megachile cincta (Megachilidae) (7.0%), Amegilla calens (Apidae) (6.2%), Xylocopa olivacea (Apidae) (5.6%), Megachile kamerunensis (Megachilidae) (4.6%), Musca domestica (Muscidae) (4.0%), Camponotus maculatus (Formicidae) (3.65%), Lasioglossum hancocki (Halictidae) (2.2%), and Pteropera carnapi (Acrididae) (1.3%) while other species were rare and represented each by less than 1.0%. Amongst these insects, two exotic Diptera (Calliphora vicina and Mu. domestica) were known as human health pests causing myiasis infections
[80-82]
. Two useful species (the exotic Eumenidae Delta sp. and the afrotropical Ascalapidae predator Ascalaphus africanus) were able to be used as biological control agents against phytophagous pest insects such as the Acrididae pest Pe. carnapi
[101, 112]
. The diversity of the recorded flower visiting insects is reminiscent of the reports from several countries including Cameroon on several plant species in market gardens
[47, 66, 67, 112-120]
. Potential pest insects with phytophagous larvae were Lepidoptera (Nymphalidae and Pieridae) and Orthoptera (Acrididae), cumulatively representing 3.1% of the total collection. Sap-feeding insects were not recorded contrary to the case in other market garden crops in Cameroon
[39]
. Our results were contrary to those reported in Se. indicum plants in Egypt, India
[123, 124]
and in other market garden plants such as cowpea, potato and eggplant in Cameroon
[66, 67]
. For illustration, it is the case in cowpea fields in Indonesia, Egypt, Nigeria
[47, 116, 125, 126]
, in cowpea fields in Cameroon
[39]
, in potato and eggplants fields in Cameroon
[39, 45, 46-53, 63-67]
where Homoptera Aphididae was higly recorded. The recorded number of species was low compared to the situation reported in Egypt where 31 insect species collected on Se. indicum plants were divided into four groups, true pollinators (Hymenoptera), other pollinators (Diptera, Coleoptera and Lepidoptera), pests (Orthoptera, Odonata, Hemiptera and Homoptera) and natural enemies (Coleoptera, Hymenoptera, Neuroptera and Dictyoptera)
[120]
. It was the same in India where Sesamum flowers attracted 24 species belonging to 17 families under eight orders in Odisha locality
[121]
and 34 insect species belonging to 18 families from four orders in Haryana locality
[124]
. The species richness of the flower-visiting insects was quite close to the observations made in cowpea fields in India where a list of 19 insect species was reported
[65]
. Similar results were reported in cowpea plantations in Cameroon where flowers were visited by insects belonging to six orders, 13 families, 19 genera and 19 species and where Coleoptera, Hemiptera and Hymenoptera were species-rich orders (five species each i.e. 26.3%) and Hemiptera was mostly abundant (40.0%) followed by Coleoptera (27.6%), Hymenoptera (21.9%), Lepidoptera (0.9%). Heteroptera and Orthoptera (0.8% respectively)
[39]
. The peculiarity of our results was the absence of five main taxa frequently recorded in market garden fields (Coleoptera, Dictyoptera, Hemiptera, Homoptera, and Odonata) certainly due to the short time period of our study (13 consecutive days from the first day of the flowering period: the week of the early blooming stage and the first week of the mid bloom stage) and probably due to a low production of the attractive scent by the blooming flowers since plants were not at their optimal flowering period. It is well known that flowers progressively appear on mature plants (reproductive period) and each blooming flower remain on the plants for ten days. The blooms do not open all at once, but gradually from the base of the stem upwards to the top of the plant
[125, 126]
. Due to the non uniform, indeterminate nature of the blooming period, the reproductive, ripening, and drying phases of the seed tend to overlap, seed lowest on the plant being mature first, even as the upper part of the plant is still flowering or has just formed seed capsules
[126]
. The duration of the early blooming stage of the mature plants (not all flowers set capsules) is one week, the duration of the mid bloom stage is four weeks (over 70% of flowers occur in first two weeks of this stage) and the duration of the last blooming stage is one week (leaves in light start to fall off), making a total of six weeks for the reproductive phase of the mature plants
[125-127]
. Even though the reproductive stage can go on for six weeks, weeks two and three produce 70-75% of the flowers and it is the most important two weeks of the cycle
[127]
. Thus, our study period would certainly only concern a few cohorts of flowers produced. In the localities of Bilone (Obala-Cameroon), natural enemies were the most recorded (Hymenoptera, and Neuroptera with one family: 92.4% of the total collection) followed by the true pollinators (Hymenoptera with six families: 91.5%) while rare taxa were other pollinators (Diptera and Lepidoptera with two families each: 6.3%), and pests (Orthoptera with one family: 1.3%), suggesting that flowers of Se. indicum were widely and frequently visited by beneficial insects (pollinators). Nevertheless, the low rate of visited flowers (20.3% during the two campaigns: 5.7% in 2022 and 14.7% in 2023) suggested either flowers did not necessarily need pollinating insects because they are hermaphrodite with facultative allogamy, producing both nectar and pollen attractive to insects
[128]
or the scarcity of associated entomofauna in neighboring sites. Moreover it was demonstrated in the locality of Bambui (North-West of Cameroon) that Se. indicum presented a mixed allogamous-autogamous reproduction regime with the predominance of autogamy
[43]
. Blooming flowers of Se. indicum produce nectar attractive to pollinator and non-pollinators, which allowed this plant species to be classified in the category of highly nectar-producing plants and weekly pollen-producing bee plants. Therefore it is necessary to preserve plants of Se. indicum and/or cultivate them not far from the hives. Consequently, bee foragers could play a positive role on geitogamy
[64]
by depositing the pollen of one flower on the stigma of another flower of the same plant. Foragers that passed from flower to flower could transport pollen from one plant to another and thus allow xenogamy by putting the pollen from one plant on the stigma of a flower belonging to another plant. Apoides are known as pollinators of Se. indicum in Egypt
[129]
and in Bambui (North-West Cameroon)
[43]
and even pollinators of cowpea in Yaounde, Maroua and Ngaoundere
[39, 44]
. Rare species included a native phytophagous Orthoptera (one family: 1.3%) known as pest for plants and two exotic Diptera (Calliphora vicina native to North America and Mu. domestica native to the Palearctic Region) known as responsible of human myiasis infections. These phytophagous insects and myiasigenic species are frequently recorded in anthropized areas
[61, 101]
. The damage caused by phytophagous insects (Coleoptera Chrysomelidae, Hymenoptera, Lepidoptera and Orthoptera) is greater on leaves and pods because at the fruiting stage, plants emit volatile compounds which attract insects (pollinators, predators and pests including phytophagous insects). The situation found in the localities of Bilone (Obala-Cameroon) in Se. indicum plots is therefore not surprising. In market garden crops (example of the reports from Ivory Coast, Egypt, Nigeria
[47, 116, 125, 126]
, and Cameroon
[39, 45, 46-53, 63-67, 130]
), aerial plant organs such as leaves, flowers and pods can be more attacked than other plant organs, depending on the high production periods. Our study is the first step in evaluating the species richness of flower visiting native and non-native insects on Se. indicum flowers. In Se. indicum, pollen is produced by the anthers which are easily accessible to the foragers, while, the nectar produced in the corollary tube, is difficult to access as already noted in Bambui (West-Cameroon)
[43]
. Plots of Se. indicum showed low species richness (richness ratio close to null), high species diversity (Shannon-Weaver index close to the maximum value), a low dominance level of a few species (Berger-Parker index inferior to the median value) and a high level of the species evenness (Pielou index close to one). Similar results are reported in ground-dwelling ants in anthropized environments
[113, 131, 132]
, in the assemblages of insects associated with potato plants
[67]
or eggplants
[66]
, in the assemblage of the floricultural insects associated with cowpea plants
[39]
. Recent reports show that the same orders and families damage chili pepper plants (Piper nigrum L.) in the locality of Penja-Cameroon
[133]
. The low diversity of the flower visiting insects was associated with a high abundance in native species, resulting in the high exploitation of resources. The exploitation of both food and nest sites was rarely achieved by non-native species (15.8% of the total species richness and 5.2% of the total abundance). These results were contrary to the reports in cowpea, egg-plant and potato fields in Cameroon
 [39, 66, 67]
. Based on the reports on the harmful activity of non-native species in the localities of introduction, they would carry out in Bilone a similar activity in sesame plots. The low representation of exotic species could be the result either of the regulation of their populations by local enemies or of unsuitable environmental conditions in the study location.
4.2. Community Structure and Functioning
Assemblage of flower-visiting insects in Bilone best fitted in 2022, the LL model with a Motomura environmental constant close to one (m=0.777). In 2023 and the pooled campaigns, settlements best fitted the LN model (Preston niche partitioning model) with in each case a Preston environmenttal constant close to one (m’=0.726 and m’=0.732 respectively). LL model reflects a community where the majority of species show moderate abundance (a community in which a reduced number of species is largely dominant, or a pioneer assemblage)
[134]
. High values of the Motomura or the Preston parameters suggest a high decay rate of abundance per rank of the species, as reported in pioneer assemblages (elementary interspecies relations with competition limited to the physical space)
[135]
. LL (niche partitioning model) is reported fitting SADs of ground-dwelling ants in France
[136]
and in Cameroon
[120]
, the dung beetles in the Southern Alps
[137]
, sand flies in Congo
[138]
, the Carabidae and Heteroptera in Finland
[139]
, grasshoppers in Cameroon
[140]
, insects associated with potato plants, eggplants and cowpea plants in Cameroon
[66, 67, 132]
. LN is reported fitting SADs of invertebrates
[138, 140, 141]
and characterizes open or less disturbed environments. It is well known that human activities in general, resulting in large deforestation, urbanization and growing cities affect ecosystem functioning and contribute to the loss of biodiversity
[142]
. A similar situation occurs in Bilone. LL niche partitioning and LN models reflect communities with moderately abundant majority of species. It is well known that nomocenosis are associations of species subject to the influence of the same factors and whose species profile is sufficiently close to be assimilated to LL or LN representation (open or more or less disturbed environments) with a strong competition between pioneer species for exploitation of available resources
[142]
.
5. Conclusion
The purpose of this study was to determine the biodiversity of the flower-visiting insects on Sesamum indicum and characterize their community structure. Specimens belonged to five orders, 12 families, 18 genera and 19 species. Hymenoptera was the most recorded (91.5% of the collected insects). Other orders were rare: Diptera (4.5%), Lepidoptera (1.8%), Neuroptera (0.9%), and Orthoptera (1.3%). Apidae was the most recorded family (42.4%) followed by Formicidae (34.1%), Megachilidae (11.6%) and other families were rare: Acrididae (1.3%), Ascalapidae (0.9%), Calliphoridae (0.5%), Eumenidae (0.7%), Halictidae (2.2%), Muscidae (4.0%), Nymphalidae (1.3%), Pieridae (0.5%), and Vespidae (0.6%). Apis mellifera (Apidae) was the most recorded species (30.6%), followed by Paratrechina longicornis (Formicidae) (12.3%), Pheidole megacephala (Formicidae) (9.4%), Myrmicaria opaciventris (Formicidae) (8.9%), Megachile cincta (Megachilidae) (7.0%), Amegilla calens (Apidae) (6.2%), Xykocopa olivacea (Apidae) (5.6%), Me. kamerunensis (Megachilidae) (4.6%), Musca domestica (Muscidae) (4.0%), Camponotus maculatus (Formicidae) (3.65%), Lasioglossum hancocki (Halictidae) ((2.2%), and Pteropera carnapi (Acrididae) (1.3%). Two exotic Diptera (Calliphora vicina and Mu. domestica) were myiasigenic species. Two useful species were recorded (the exotic Eumenidae Delta sp. and the Ascalapidae predator Ascalaphus africanus). Assemblages showed low species richness, high species diversity, a low dominance of a few species and the community was highly even. The sampling success was maximal. The number of simply abundant species was close to the number of co-dominants. Overall, insects exhibited in 2022 and 2023, a positive net association in presence/absence data. A few negative correlations (mutual repulsion) and several positive correlations (mutual tolerance) were noted. Calliphora vicina (Calliphoridae) was negatively correlated with Ap. mellifera (Apidae). Positive correlation was recorded in several combinations and the community functioned on the base of niche partritionning models (LL in 2022, LN in 2023 and the pooled years), suggesting a amore or less disturbed environment with a strong competition between pioneer species for the available resources.
Abbreviations

Ac. acerata

Acraea acerata Hewitson. 1874

ACE

Abundance Coverage-based Estimator

AIC

Akaike Information Criteria

Am. calens

Amegilla calens (Lepeletier De Saint-Fargeau.1841)

Ap. mellifera adansonii

Apis mellifera adansonii Latreille. 1804

As. africanus

Ascalaphus africanus (McLachlan. 1871)

Bi. dorothea

Bicyclus dorothea (Cramer. 1779)

BC

Bray-Curtis Index

BIC

Bayesian Information Criteria

BS

Broken-Stick Theoretical Model

Ca. maculatus

Camponotus maculatus (Fabricius. 1782)

Cl. vicina

Calliphora vicina Robineau-Desvoidy. 1830

Ct. florella

Catopsilia florella (Fabricius. 1775)

CRC

Central Regional Council

FAOSTAT

Food and Agricultural Organization Statistics

GBIF

Global Biodiversity Information Facility

GM

Geometric Theoretical Model

IRAD/ARID

Institut de Recherche Agricole Pour le Développement/Agricultural Research Institute for Development

ICE

Incidence Coverage-based Estimatorç

ITIS

Integrated Taxonomic Information System

OHIAM

Obala Higher Institute of Agriculture and Management

La. hancocki

Lasioglossum (Ipomalictus) hancocki (Cockerell 1945)

LL

Loglinear Theoretical Model

LN

Lognormal Theoretical Model

Me. kamerunensis

Megachile (Chalicodoma) kamerunensis Friese. 1922

Me. cincta

Megachile (Chalicodoma) cincta (Fabricius. 1781)

Mu. domestica

Musca domestica Linnaeus. 1758

My. opacivenytrtis

Myrmicaria opaciventris Emery. 1893

Pa. longicornis

Paratrechina longicornis (Latreille. 1802)

Pe. carnapi

Pteropera carnapi Ramme. 1929

Ph. megacephala

Pheidole megacephala (Fabricius. 1793)

POWO

Plant of the World Online

SAD

Species Abundance Distribution

sp.

Undetermined Species

Se. alatum

Sesamum alatum Thonn.

Se. indicum

Sesamum indicum L. (1753)

So. tuberosum

Solanum tuberosum L., 1753

SPSS

Statistical Package for the Social Sciences

Sy. conuta

Synagris conuta (Linnaeus. 1758)

VR

Variance Ratio

Xy. olivacea

Xylocopa olivacea (Fabricius 1778)

Z

Zipf Model

ZM

Zipf-Mandelbrot Model
Acknowledgments
The authors acknowledge the Cameroonian Ministry of Higher Education for providing funds through the research support program. They thank the elders of the Laboratory of Applied Zoology of University of Ngaoundere for assistance in the insect identification and the manuscript preparation.
Author Contributions
Pharaon Auguste Mbianda: Conceptualization, Data curation, Formal Analysis, Investigation, Methodology, Writing - original draft, Writing - review & editing
Moukhtar Mohammadou: Formal Analysis, Software, Writing - original draft
Taïmanga: Formal Analysis, Software, Writing - original draft
Andrea Sarah Kenne Toukem: Formal Analysis, Software, Writing - original draft
Sedrick Junior Tsekane: Formal Analysis, Software, Writing - original draft
Alice Virginie Tchiaze Ifoue: Formal Analysis, Software, Writing - original draft
Xavier Arthur Nyoumi Ongolo: Formal Analysis, Software, Writing - original draft
Dounia Dounia: Formal Analysis, Software, Writing - original draft
Nadine Esther Otiobo Atibita: Formal Analysis, Software, Writing - original draft
Chantal Douka: Formal Analysis, Software, Writing - original draft
Joseph Blaise Pando: Formal Analysis, Software, Writ ing - original draft
Fernand-Nestor Tchuenguem Fohouo: Conceptualization, Data curation, Formal Analysis, Methodology, Software, Writing - original draft, Writing - review & editing
Martin Kenne: Conceptualization, Data curation, Formal Analysis, Funding acquisition, Project administration, Resources, Software, Supervision, Validation, Writing - original draft, Writing - review & editing Data
Data Availability Statement
The data is available from the corresponding author upon reasonable request.
Conflicts of Interest
The authors declare no conflicts of interest.
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    Mbianda, A. P., Mohammadou, M., Taïmanga, Kenne, A. S., Tsekane, S. J., et al. (2025). Diversity, Abundance and the Community Structure of the Flower-Visiting Insects on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) in Bilone (Obala-Cameroon). American Journal of Entomology, 9(1), 28-54. https://doi.org/10.11648/j.aje.20250901.14

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    Mbianda, A. P.; Mohammadou, M.; Taïmanga; Kenne, A. S.; Tsekane, S. J., et al. Diversity, Abundance and the Community Structure of the Flower-Visiting Insects on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) in Bilone (Obala-Cameroon). Am. J. Entomol. 2025, 9(1), 28-54. doi: 10.11648/j.aje.20250901.14

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    Mbianda AP, Mohammadou M, Taïmanga, Kenne AS, Tsekane SJ, et al. Diversity, Abundance and the Community Structure of the Flower-Visiting Insects on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) in Bilone (Obala-Cameroon). Am J Entomol. 2025;9(1):28-54. doi: 10.11648/j.aje.20250901.14

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  • @article{10.11648/j.aje.20250901.14,
  author = {Auguste Pharaon Mbianda and Moukhtar Mohammadou and Taïmanga and Andrea Sarah Kenne and Sedrick Junior Tsekane and Alice Virginie Tchiaze Ifoue and Xavier Arthur Nyoumi Ongolo and Dounia Dounia and Nadine Esther Otiobo Atibita and Chantal Douka and Joseph Blaise Pando and Fernand-Nestor Tchuenguem Fohouo and Martin Kenne},
  title = {Diversity, Abundance and the Community Structure of the Flower-Visiting Insects on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) in Bilone (Obala-Cameroon)
},
  journal = {American Journal of Entomology},
  volume = {9},
  number = {1},
  pages = {28-54},
  doi = {10.11648/j.aje.20250901.14},
  url = {https://doi.org/10.11648/j.aje.20250901.14},
  eprint = {https://article.sciencepublishinggroup.com/pdf/10.11648.j.aje.20250901.14},
  abstract = {In order to identify flower-visiting insects on sesame plants and characterize the community structure, ecological survey was conducted in Bilone agroecological farm in 2022 and 2023, in 15 experimental plots (6x5.5 m each) each year, created in a 1,600 m² area. Insects were captured, stored in papillotes (Lepidoptera) or in vials containing 70° alcohol (other adults) and identified at the species level in laboratory. A total of 1,703 specimens were captured. They belonged to five orders, 12 families, 18 genera and 19 species. Hymenoptera was mostly collected order (91.5%) followed by Diptera (4.5%), Lepidoptera (1.8%), Neuroptera (0.9%) and Orthoptera (1.3%). Apidae was the most collected family (42.4%) followed by Formicidae (34.1%), Megachilidae (11.6%) while other families were rare: Acrididae (1.3%), Ascalapidae (0.9%), Calliphoridae (0.5%), Eumenidae (0.7%), Halictidae (2.2%), Muscidae (4.0%), Nymphalidae (1.3%), Pieridae (0.5%), and Vespidae (0.6%). Apis mellifera adansonii (Apidae: 30.6%) was the most recorded species, followed by Paratrechina longicornis (Formicidae: 12.3%), Pheidole megacephala (Formicidae: 9.4%), Myrmicanioa opaciventris (Formicidae: 8.9%), Megachile cincta (Megachilidae: 7.0%), Amegilla calens (Apidae: 6.2%), Xylocopa olivacea (Apidae: 5.6%), Megachile kamerunensis (Megachilidae: 4.6%), Musca domestica (Diptera: 4.0%), Camponotus maculatus (Formicidae: 3.65%), Lasioglossum hancocki (Halictidae: 2.2%), and Pteropera carnapi (Acrididae: 1.3%). Calliphora vicina (Calliphoridae) was recorded exclusively in 2022. Two exotic Diptera (Cl. vicina and Mu. domerstica) were myiasigenic species. The exotic Eumenidae Delta sp. and the afrotropical predator Ascalaphus africanus (Ascalapidae) were recorded as well as the phytophagous Acrididae Pe. carnapi. Potential pests (Nymphalidae, Pieridae and Acrididae) cumulatively represented 3.1% of the collection. The community was highly diversed and lowly dominated by a few species. Ca. maculatus was simply abundant in 2023. Amegilla calens, Ap. mellifera adansonii, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Ph. megacephala and Xy. olivacea were simply abundant. Amegilla calens and Me. cincta, were co-dominants in 2022. Ca. maculatus and Me. kamerunensis were co-dominants in 2023. Apis mellifera adansonii, Pa. longicornis, Ph. megacephala, My. opaciventris and Xy. olivacea were co-dominants in each year. Ca. maculates and Cl. vicina were rare in 2022. Bicyclus dorothea (Nymphalidae), Delta sp. and La. hancocki were rare in 2023. Acraea acerata (Nymphalidae), Ascalaphus africanus (Ascalapidae), Catopsilia florella (Pieridae), Pteropera carnapi (Acrididae) and Synagris conuta (Vespidae) were rare. High value of Motomura constant (m=0.777 in 2022) and Preston constant (m=0.726 in 2023) suggested least evolved pioneer assemblages with species competition limited to the physical space. Overall, flower visiting insects exhibited a global positive net association.
},
 year = {2025}
}

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  • TY  - JOUR
T1  - Diversity, Abundance and the Community Structure of the Flower-Visiting Insects on Sesamum indicum L. (1753) (Scrophulariales: Pedaliaceae) in Bilone (Obala-Cameroon)

AU  - Auguste Pharaon Mbianda
AU  - Moukhtar Mohammadou
AU  - Taïmanga
AU  - Andrea Sarah Kenne
AU  - Sedrick Junior Tsekane
AU  - Alice Virginie Tchiaze Ifoue
AU  - Xavier Arthur Nyoumi Ongolo
AU  - Dounia Dounia
AU  - Nadine Esther Otiobo Atibita
AU  - Chantal Douka
AU  - Joseph Blaise Pando
AU  - Fernand-Nestor Tchuenguem Fohouo
AU  - Martin Kenne
Y1  - 2025/01/21
PY  - 2025
N1  - https://doi.org/10.11648/j.aje.20250901.14
DO  - 10.11648/j.aje.20250901.14
T2  - American Journal of Entomology
JF  - American Journal of Entomology
JO  - American Journal of Entomology
SP  - 28
EP  - 54
PB  - Science Publishing Group
SN  - 2640-0537
UR  - https://doi.org/10.11648/j.aje.20250901.14
AB  - In order to identify flower-visiting insects on sesame plants and characterize the community structure, ecological survey was conducted in Bilone agroecological farm in 2022 and 2023, in 15 experimental plots (6x5.5 m each) each year, created in a 1,600 m² area. Insects were captured, stored in papillotes (Lepidoptera) or in vials containing 70° alcohol (other adults) and identified at the species level in laboratory. A total of 1,703 specimens were captured. They belonged to five orders, 12 families, 18 genera and 19 species. Hymenoptera was mostly collected order (91.5%) followed by Diptera (4.5%), Lepidoptera (1.8%), Neuroptera (0.9%) and Orthoptera (1.3%). Apidae was the most collected family (42.4%) followed by Formicidae (34.1%), Megachilidae (11.6%) while other families were rare: Acrididae (1.3%), Ascalapidae (0.9%), Calliphoridae (0.5%), Eumenidae (0.7%), Halictidae (2.2%), Muscidae (4.0%), Nymphalidae (1.3%), Pieridae (0.5%), and Vespidae (0.6%). Apis mellifera adansonii (Apidae: 30.6%) was the most recorded species, followed by Paratrechina longicornis (Formicidae: 12.3%), Pheidole megacephala (Formicidae: 9.4%), Myrmicanioa opaciventris (Formicidae: 8.9%), Megachile cincta (Megachilidae: 7.0%), Amegilla calens (Apidae: 6.2%), Xylocopa olivacea (Apidae: 5.6%), Megachile kamerunensis (Megachilidae: 4.6%), Musca domestica (Diptera: 4.0%), Camponotus maculatus (Formicidae: 3.65%), Lasioglossum hancocki (Halictidae: 2.2%), and Pteropera carnapi (Acrididae: 1.3%). Calliphora vicina (Calliphoridae) was recorded exclusively in 2022. Two exotic Diptera (Cl. vicina and Mu. domerstica) were myiasigenic species. The exotic Eumenidae Delta sp. and the afrotropical predator Ascalaphus africanus (Ascalapidae) were recorded as well as the phytophagous Acrididae Pe. carnapi. Potential pests (Nymphalidae, Pieridae and Acrididae) cumulatively represented 3.1% of the collection. The community was highly diversed and lowly dominated by a few species. Ca. maculatus was simply abundant in 2023. Amegilla calens, Ap. mellifera adansonii, Me. cincta, Me. kamerunensis, Mu. domestica, My. opaciventris, Pa. longicornis, Ph. megacephala and Xy. olivacea were simply abundant. Amegilla calens and Me. cincta, were co-dominants in 2022. Ca. maculatus and Me. kamerunensis were co-dominants in 2023. Apis mellifera adansonii, Pa. longicornis, Ph. megacephala, My. opaciventris and Xy. olivacea were co-dominants in each year. Ca. maculates and Cl. vicina were rare in 2022. Bicyclus dorothea (Nymphalidae), Delta sp. and La. hancocki were rare in 2023. Acraea acerata (Nymphalidae), Ascalaphus africanus (Ascalapidae), Catopsilia florella (Pieridae), Pteropera carnapi (Acrididae) and Synagris conuta (Vespidae) were rare. High value of Motomura constant (m=0.777 in 2022) and Preston constant (m=0.726 in 2023) suggested least evolved pioneer assemblages with species competition limited to the physical space. Overall, flower visiting insects exhibited a global positive net association.

VL  - 9
IS  - 1
ER  -

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Author Information

  • Auguste Pharaon Mbianda

    Department of Plant Biology, University of Douala, Douala, Cameroon

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0009-0004-8595-9700

  • Moukhtar Mohammadou

    Department of Biology and Physiology of Animal Organisms, University of Douala, Douala, Cameroon

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0009-0008-8986-4150

  • Taïmanga

    Department of Agronomy, University of Douala, Douala, Cameroon

    Research Fields: Botanical insecticides, Bees, Biological agriculture, Yield agricultural products, Cowpea production, Pollinization

    Contact Email

    http://orcid.org/0009-0002-3650-2638

  • Andrea Sarah Kenne

    Department of Biology and Physiology of Animal Organisms, University of Douala, Douala, Cameroon

    Research Fields: Water toxicology, Waterborne diseases, Water quality of life, Biostatistics, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0009-0008-6261-3984

  • Sedrick Junior Tsekane

    Department of Biology and Physiology of Animal Organisms, University of Douala, Douala, Cameroon

    Research Fields: Applied zoology, Quality of life and biostatistics, Wildlife Protection, Control of protected areas, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0000-0002-7225-8689

  • Alice Virginie Tchiaze Ifoue

    Department of Plant Biology, University of Douala, Douala, Cameroon

    Research Fields: Applied entomology, Insects biology, biostatistics, pollinators, Plant physiology, Soil fertilization

    Contact Email

    http://orcid.org/0009-0006-0015-380X

  • Xavier Arthur Nyoumi Ongolo

    Department of Agronomy, University of Dschang, Obala, Cameroon

    Research Fields: Applied entomology, Insects biology, Pollinators, Seeds production, Agriculture, Plant variety amelioration.

    Contact Email

    http://orcid.org/0009-0007-7416-8254-

  • Dounia Dounia

    Higher Teacher Training College, University of Yaounde 1, Yaounde, Cameroun

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0000-0002-9342-8254

  • Nadine Esther Otiobo Atibita

    Department of Biology and Physiology of Animal Organisms, University of Douala, Douala, Cameroon

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0000-0002-2554-0869

  • Chantal Douka

    Higher Teacher Training College, University of Yaounde 1, Yaounde, Cameroun

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0000-0002-3657-1775

  • Joseph Blaise Pando

    Higher Teacher Training College, University of Maroua, Maroua, Cameroun

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0009-0001-2243-2002

  • Fernand-Nestor Tchuenguem Fohouo

    Department of Biological Sciences, University of Ngaoundere, Ngaoundere, Cameroon

    Research Fields: Applied entomology, Insects biology, biostatistics and pollinators, Apiculture, Insects-plants interactions, Animal Ethology, Animal Ecology

    Contact Email

    http://orcid.org/0000-0002-1098-9866

  • Martin Kenne

    Department of Biology and Physiology of Animal Organisms, University of Douala, Douala, Cameroon

    Research Fields: Biostatistics and Biology of the Animal Populations, Entomology and Myrmecology, Animal Ethology, Animal Ecology and sociobiology, Applied entomology and plant protection, Biological control of pest insects

    Contact Email

    http://orcid.org/0000-0002-2104-7073

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  • Abstract
  • Keywords

  • Document Sections

    1. 1. Introduction
    2. 2. Material and Methods
    3. 3. Results
    4. 4. Discussion
    5. 5. Conclusion
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  • Abbreviations
  • Acknowledgments
  • Author Contributions
  • Data Availability Statement
  • Conflicts of Interest
  • References
  • Cite This Article
  • Author Information

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