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ORIGINAL ARTICLE
A study on fruit fly host range reveals the low infestation rate
of Bactrocera dorsalis (Tephritidae) in Mayotte
Laura Moquet
1
| Tim Dupin
1
| Louis Maigné
1
| Joel Huat
2
|
Thomas Chesneau
3
| Hélène Delatte
4
1
CIRAD, UMR PVBMT, Saint-Pierre, Reunion
2
CIRAD, HORTSYS, Saint-Pierre, Reunion
3
Etablissement Public National, Coconi,
Mayotte
4
CIRAD, UMR PVBMT, Antananarivo,
Madagascar
Correspondence
Laura Moquet, Pôle de Protection des Plantes,
7 Chemin de l’Irat, 97410 Saint-Pierre,
Réunion.
Email: laura.moquet@cirad.fr
Funding information
European Funds for Rural Development
(EFRD); INNOVEG project (Network for
Innovation and Transfer in Agriculture, RITA);
European Agricultural Fund for Rural
Development; Centre de Coopération
Internationale en Recherche Agronomique
pour le Développement; French Ministry of
Agriculture (Ministère de l’Agriculture, de
l’Agroalimentaire et de la Forêt); Région
Réunion
Abstract
1. The oriental fruit fly (Bactrocera dorsalis) is one of the world’s most invasive and
polyphagous fruit pests. It causes severe damage throughout its range and can dev-
astate the entire fruit harvest in unprotected orchards. In 2007, B. dorsalis was
detected in Mayotte, where it now ranks ninth on the list of fruit fly species of eco-
nomic importance. This tropical island is a good study area to analyse the host
range of B. dorsalis and its interactions with other resident fruit fly species.
2. Two field campaigns were carried out from 2012 to 2014 and from 2019 to 2021.
We collected fruit from all over the island in cultivated and non-cultivated areas
and compared the infestation rates between the two periods.
3. We detected six fruit fly species, including the common species Dacus ciliatus, Neo-
ceratitis cyanescens, Ceratitis capitata, B. dorsalis and two rarer species, Dacus etien-
nellus and Trirhithrum nigerrimum.
4. The most surprising result was the low occurrence of B. dorsalis, with only seven
host plant species identified out of a total of 84 plant species. Infestation rates
were low for these host plant species, even in the case of mango (11.71 flies/kg)
and Indian almond (0.97 fly/kg), which are considered to be major host plants of
B. dorsalis.
5. Bactrocera dorsalis seems to have a lower impact in Mayotte than in other parts of
the world. We discuss the possible causes of the weak infestation rates observed,
which could provide the key to regulating the species on the island.
KEYWORDS
biological invasions, food-web network, Indian Ocean, infestation rates, oriental fruit fly
INTRODUCTION
The oriental fruit fly, Bactrocera dorsalis (Hendel, 1912), (Diptera,
Tephritidae), endemic to the Indo-Asian region, is one of the world’s
most invasive and polyphagous pests of fruits and vegetables
(White & Elson-Harris, 1992). This fruit fly species has the widest host
range within the Bactrocera genus and has been recorded on more
than 500 host plants (Allwood et al., 1999; Clarke et al., 2005; Liquido
et al., 2015). Its extensive host range allows it to sustain populations
both spatially and temporally all year round. Bactrocera dorsalis causes
crop damage, which can have a major economic impact and lead to
the loss of export markets. For example, in some cultivated species,
such as mango and guava, up to 100% of fruit can be infested (Badii
et al., 2015). Bactrocera dorsalis is highly competitive and can displace
Received: 4 July 2023 Accepted: 18 January 2024
DOI: 10.1111/afe.12614
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction in any
medium, provided the original work is properly cited and is not used for commercial purposes.
© 2024 The Authors. Agricultural and Forest Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society.
Agr Forest Entomol. 2024;1–13. wileyonlinelibrary.com/journal/afe 1
ecological niches of pre-established tephritid species, such as Ceratitis
sp. or other Bactrocera sp. (Sauders, 1841) (Duyck et al., 2004; Ekesi
et al., 2016; Hassani, 2017; Moquet et al., 2021; Mwatawala
et al., 2006; Rasolofoarivao et al., 2021). This species often becomes
the dominant generalist fruit fly species in invaded locations. This has
occurred on islands in the Indian Ocean, such as Madagascar,
Mauritius and La Réunion (Moquet et al., 2021; Rasolofoarivao
et al., 2021; Sookar et al., 2021).
In Mayotte, B. dorsalis was first recorded in 2007 (De Meyer
et al., 2010,2012), but its host range, infestation rate and interactions
with other species have yet to be investigated. In Mayotte, nine species
of Tephritidae were recorded, including one endemic to the archipel-
ago, Dacus etiennellus Munro, 1984 (De Meyer et al., 2012). Besides
B. dorsalis, the known invasive fruit fly pests on the island include Dacus
ciliatus Loew, 1862, which generally infests Cucurbitaceae (Ryckewaert
et al., 2010); Neoceratitis cyanescens (Bezzi, 1923), which is found on
Solanaceae; and the polyphagous species, Ceratitis capitata
(Wiedemann, 1824) (De Meyer et al., 2012; Franck & Delatte, 2020).
Other fruit fly species have been reported in Mayotte, including Dacus
bivittatus (Bigot,1858) and Dacus vertebratus Bezzi, 1908, on Cucurbita-
ceae; and Trirhithrum nigerrimum (Bezzi, 1913) and Ceratitis malgassa
(Munro, 1939), which attack a wide variety of different host plant fami-
lies (De Meyer et al., 2012; Rasolofoarivao et al., 2021).
Agriculture is one of the principal activities in Mayotte. It provides
a livelihood for around 60,000 people, a third of the population
(DAAF Mayotte, 2021). Local agriculture supplies about 80% of the
fresh fruits and vegetables consumed on the island (SISE/DAAF
Mayotte, 2017). Most farmers use traditional agricultural and food
systems called ‘Mahorais gardens’. These are small-scale agroforestry
systems, which are multilayered and include various cultural associa-
tions (DAAF Mayotte, 2016). These systems often include fruit trees
(mango, orange, etc.), fruit and leaf vegetables, roots and tubers, aro-
matic plants (vanilla, ylang-ylang, etc.), forage plants and other food
crops.
Our aim was to identify the host range of fruit flies of economic
importance in Mayotte and to determine the diversity of parasitoids
that attack tephritid species. We focused particular attention on
B. dorsalis, the most recent invasive species recorded on the island. To
study the changes in fruit fly infestation over time, we compared
infestation rates between two periods: 5–7 years and 12–14 years
after the invasion of B. dorsalis.
MATERIALS AND METHODS
Study region
Mayotte is part of the Comoros Archipelago. It is located in the
Mozambique Channel in the Indian Ocean, about 300 kilometres
northwest of Madagascar. Mayotte is split into two main islands,
Grande-Terre (or Mahoré) and Petite-Terre (or Pamandzi). Mayotte’s
climate is tropical and humid with two distinct seasons: a warm humid
rainy season from December to March (temperature ranges from
24 to 32C with 70%–95% humidity), and a cooler dry season from
June to September (temperature ranges from 20 to 28C with 61%–
90% humidity) (Météo-France, 2022; Appendix S1, Figure S1). These
abiotic factors are highly suitable for the development of tropical fruit
flies, especially B. dorsalis (De Villiers et al., 2015).
Mayotte has very diverse landscapes, and trees constitute a major
component, alternating between forests and ‘agroforestry’systems.
The dense forest cover represents approximately 13,730 ha or 36.7%
of the island’s total surface area, whereas the primary forests repre-
sent only 5% (Lartigue & Boisseaux, 2020; Appendix S1, Figure S2).
Although some fruits were collected in natural forests (Mimusops
comoriensis, Ficus sycomorus, etc.), our study was largely conducted in
agroforestry systems, where the dominant woody species are jackfruit
(Artocarpus heterophyllus), breadfruit (Artocarpus altilis), African tulip
tree (Spathodea campanulata), mango (Mangifera indica) and coconut
palm (Cocos nucifera). There are two main types of agroforestry sys-
tems: the first combines crops with a more or less dense cover of fruit
species and the second combines food or cash crops (vanilla, turmeric,
coffee, etc.) with a cover of forest species (Lartigue &
Boisseaux, 2020). Pure vegetable systems are less common, particu-
larly located in the central area of the island. Vegetables are generally
grown in the dry season, and the main crops include peppers, toma-
toes, cucumbers, zucchini and aubergine.
Sampling
We collected fruit during two separate periods from 2012 to 2014
and from 2019 to 2021. Field campaigns covered the two main islands
(Grande-Terre and Petite Terre, Figure 1). When possible, samples
were collected every week from January 2012 to March 2013, from
April 2019 to July 2020, and from March 2021 to July 2021. We
scouted the island to collect any soft-skinned fruit. Additional sam-
plings were conducted in July 2013, January 2014 and December
2020. We took samples of any fleshy fruit species, cultivated or wild.
In cultivated areas, we favoured non-treated crops. Fruit was ran-
domly collected from trees or from the ground when ripe, which is
when they are most susceptible to fruit fly infestation. Cucurbit fruit
was collected in the early stage (when the fruit was small and its skin
soft), whereas most other fruit was collected when mature. We col-
lected fruit from 28 plant species during the first period and from
74 plant species during the second period (Table 1). Overall, 18 species
were the same for the two sampling periods. A total of 1307 fruits
were collected between 2012 and 2014, and 6673 fruits between
2019 and 2021. We focused particularly on host plant species of
socioeconomic importance: (i) mango (Mangifera indica) from the local
variety ‘Nounou’and from spontaneous germination; and cultivated
Rutaceae (Citrus reticulata and Citrus sinensis), as potential host plants
for the generalist species, B. dorsalis; (ii) tomato (Lycopersicon esculen-
tum), as a potential host plant of the fruit fly that specialises in Solana-
ceae, Neoceratitis cyanescens; and (iii) cultivated Cucurbitaceae
(Cucumis sativus and Cucurbita pepo), as potential host plants for Dacus
ciliatus and D. etiennellus. For mango, two types of samples were
2MOQUET ET AL.
collected: small immature fruit that had just fallen from the tree
(<50 g), and large ripe fruit still on the tree or fleshy fallen (>50 g).
Incubation of fruit samples
In the laboratory, fruits were weighed, placed in plastic boxes with
sand as a pupation substrate and covered with fine mesh cloth. We
put fruit samples in a maturation room (28 ± 2C, 80 ± 10% RH) until
pupation. From 2012 to 2014, fruits from the same species collected
on the same date, within the same site, were counted and grouped for
weighing and incubation. Since 2019, fruit was weighed and incu-
bated individually. Fruit samples were inspected each week for
3 weeks, and the sand was sifted for pupae. At the end of the incuba-
tion period, fruit was opened to check for larvae and pupae. Pupae
were kept in a climate room in plastic boxes until fruit flies or parasit-
oids emerged. Individuals were then sexed and taxonomically identi-
fied to species level using morphological characteristics. We
calculated the fruit fly infestation rate as the number of individuals
per kilogram of collected fruit. Proportions of infested fruits (fruits
that allow the development of at least one pupa) were only calculated
for the period 2019–2021, when emergence from individual fruits
was recorded. We calculated the parasitism rate as the number of
emerged parasitoids over the number of emerged fruit flies.
Comparison of Bactrocera dorsalis infestation between
the two periods
We compared the infestation rate of B. dorsalis for the two sampling
periods: 2012–2014 and 2019–2021. We considered the host plant
species for which we had enough samples from both collection
periods: C. reticulata, C. sinensis, M. indica and Terminalia catappa.We
used generalised linear mixed effect models (GLMMs), with the period
as a fixed effect and plant species as a random factor. Analyses were
performed using the R software (R Core Team, 2022).
Network
The function ‘networklevel’of the ‘bipartite’package (Dormann
et al., 2008,2009) was used to determine indices describing networks
FIGURE 1 Sampling sites for the main plant species collected between (a) 2012 and 2014, and (b) 2019 and 2021.
FRUIT FLY INFESTATION RATE IN MAYOTTE 3
TABLE 1 Collected plant species in Mayotte during the period 2012–2014 and 2019–2021 to study the Tephritidae host range.
Family Species name English name
NTotal weight (g) Pupae/kg
2012–
2014
2019–
2021
2012–
2014
2019–
2021
2012–
2014
2019–
2021
Anacardiaceae Mangifera indica
a
Mango 57 882 16,149 62,065 10.5 18.1
Mango small fruits
(<50 g)
5 475 15,978 9471 251.5 16.0
Mango large fruits
(>50 g)
52 407 171 52,594 2.5 18.5
Spondias dulcis
a
Golden apple 0 47 0 4124 _ 0
Annonaceae Annona cherimola
a
Cherimoya 0 11 0 142 _ 0
Annona muricata
a
Soursop 0 12 0 6398 _ 2.5
Annona reticulata
a
Custard apple 0 21 0 3510 _ 0
Cananga odorata
a
Ylang-ylang 0 45 0 212 _ 0
Apocynaceae Petchia erythrocarpa 0 9 0 5.5 _ 0
Saba comorensis
a
Bungo fruit 0 45 0 2279 _ 0
Cascabela thevetia
a
Yellow oleander 0 76 0 932 _ 0
Boraginaceae Cordia myxa
a
Assyrian plum 0 62 0 137 _ 0
Ehretia cymosa 0 312 0 121 _ 0
Caricaceae Carica papaya
a
Papaya 0 69 0 96,812 _ 0
Combretaceae Terminalia catappa
a
Tropical almond 41 102 1558 3594 3.2 3.6
Cucurbitaceae Citrullus lanatus
a
Water melon 18 0 2041 0 71.5 _
Cucumis anguria West indian gherkin 12 0 888 0 46.2 _
Cucumis melo
a
Muskmelon 14 1 3507 701 36.5 0
Cucumis sativus
a
Cucumber 151 16 3463 4044 209.9 9.39
Cucurbita moschata
a
Butternut squash 34 41 1056 466 371.2 219.1
Cucurbita pepo
a
Zucchini 134 4 3256 2574 401.1 11.3
Lagenaria siceraria
a
Calabash 7 0 56 0 857.1 _
Luffa acutangula
a
Angled luffa 20 0 343 0 113.7 _
Momordica charantia
a
Bitter squash 5 30 30 731 0 28.7
Sechium edule
a
Chayotte 25 0 4222 0 57.8 _
Euphorbiaceae Jatropha curcas Physic nut 0 68 0 866 _ 0
Ricinus communis Castor oil tree 0 32 0 38 _ 0
Fabaceae Canavalia sp. 0 4 0 121 _ 0
Cassia fistula Golden tree 0 15 0 430 _ 0
Pithecellobium dulce Manila tamarind 0 15 0 194 _ 0
Vigna sp. 0 131 0 31 _ 0
Lauraceae Cinnamomum verum Cinnamon 0 30 0 15 _ 0
Litsea glutinosa Brown bollywood 0 379 0 145 _ 0
Persea americana
a
Avocado tree 2 15 795 38 0 0
Malvaceae Cola sp.
a
0 30 0 1362 _ 0
Theobroma cacao Cacao tree 0 4 0 1009 _ 0
Melastomataceae Clidemia hirta Koster’s curse 0 165 0 93.5 _ 0
Monimiaceae Tambourissa
leptophylla
0 1 0 603 _ 0
Moraceae Artocarpus altilis
a
Breadfruit 0 5 0 4931 _ 0
Artocarpus
heterophyllus
a
Jackfruit 0 20 0 5677 _ 0
Morus kagayamae Japanese mulberry 0 15 0 35 _ 0
Ficus benjamina 045 021_ 0
Ficus sycomorus
a
Sycamore fig 0 104 0 794 _ 13.9
4MOQUET ET AL.
TABLE 1 (Continued)
Family Species name English name
NTotal weight (g) Pupae/kg
2012–
2014
2019–
2021
2012–
2014
2019–
2021
2012–
2014
2019–
2021
Muntingiaceae Muntingia calabura Jamaica cherry 0 15 0 35.5 _ 0
Musaceae Musa sp.
a
Banana tree 0 12 0 867 _ 0
Myrtaceae Eugenia uniflora
a
Pitanga 0 27 0 188 _ 0
Psidium cattleianum
a
Purple guava 0 15 0 91 _ 0
Psidium guajava
a
Guava 6 58 93 2931 0 36.5
Syzygium malaccense
a
Malay rose apple 0 45 0 1519 _ 0
Oxalidaceae Averrhoa bilimbi
a
Bilimbi 0 131 0 3028 _ 0
Averrhoa carambola
a
Carambola 2 61 45 2024 0 0.3
Passifloraceae Passiflora foetida Bush passion fruit 0 16 0 19 _ 0
Passiflora suberosa Cork passion fruit 0 30 0 21 _ 736.8
Petiveriaceae Rivinia humilis Bloodberry 0 45 0 8 _ 0
Phyllantaceae Flueggea virosa White berry bush 0 195 0 23 _ 0
Rhamnaceae Colubrina asiatica Asian nakedwood 0 15 0 266 _ 0
Rosaceae Eriobotrya japonica
a
Japanese medlar 0 15 0 97 _ 0
Rubus alceifolius Giant bramble 0 15 0 40.5 _ 0
Rubiaceae Coffea sp.
a
Coffee 0 171 0 671 _ 0
Morinda citrifolia Indian mulberry 0 27 0 2985 _ 0
Rutaceae Citrus aurantifolia
a
Lime 0 48 0 1899 _ 0
Citrus hystrix Kaffir lime 0 26 0 1342 _ 0
Citrus limon
a
Lemon 32 66 2210 4500 0 0
Citrus medica 0 4 0 664 _ 0
Citrus reticulata
a
Tangerine 83 109 8360 10,783 1.7 1.8
Citrus sinensis
a
Orange 218 716 31,041 97,814 1.4 0.1
Citrus sinensis x Citrus
limon
18 0 2560 0 0 _
Citrus x paradisi
a
Pomelo 1 0 200 0 0 _
Citrus x tangelo
a
Tangelo 1 0 65 0 0 _
Sapindaceae Allophylus bicruris 0 111 0 19 _ 0
Sapotaceae Mimusops comorensis 0 90 0 695 _ 214.6
Mimusops coriacea
a
0 5 0 189 _ 0
Solanaceae Capsicum sp.
a
Pepper 35 269 159 466 150.7 195.3
Lycopersicon
esculentum
a
Tomato 153 126 413 2902 283.3 85.8
Solanum aethiopicum
a
Ethiopian nightshade 18 0 148 0 527 _
Solanum melongena
a
Eggplant 2 85 55 2211 145.5 28
Solanum nigrum
a
Blackberry
nightshade
38 95 5 20 2600 0
Solanum
seaforthianum
a
Brazilian nightshade 0 66 0 40 _ 0
Solanum sisymbriifolium Sticky nightshade 0 15 0 236 _ 0
Solanum torvum
a
Turkey berry 37 270 43 297 0 0
Verbenaceae Duranta repens Golden dewberry 0 232 0 113 _ 0
Lantana camara Lantana 0 75 0 20 _ 0
Lantana triofolia Threeleaf
shrubverbena
045 0 8_ 0
(Continues)
FRUIT FLY INFESTATION RATE IN MAYOTTE 5
(connectance, links per species, number of compartments, cluster
coefficient, nestedness and H2’) for each period studied. We tested if
the network presented specific patterns or corresponded to what was
forecast in the absence of a structuring mechanism, by comparing the
indices observed to indices of random webs. Thus, we performed a
t-test using the function ‘null.t.test’, where the random matrices were
based on the function ‘r2dtable’(N=1000). In addition, as explained
by Dormann (2022), we also built 1000 null models (function ‘nullmo-
del’) and computed indices for each one. We graphically compared
the distribution of null-model index values to our observed index
TABLE 1 (Continued)
Family Species name English name
NTotal weight (g) Pupae/kg
2012–
2014
2019–
2021
2012–
2014
2019–
2021
2012–
2014
2019–
2021
Premma serratifolia Headache tree 0 115 0 10 _ 0
Vitaceae Leea guinensis Hawaiian holy 0 101 0 39 _ 0
Note:N, Number of collected samples; _, No data.
a
Host plants of B. dorsalis according to Badii et al., 2015; Franck & Delatte, 2020; Goergen et al., 2011; Moquet et al., 2021; Mwatawala et al., 2006;
Ndiaye et al., 2012; Rattanapun, 2009; Rwomushana et al., 2008; Vargas et al., 2007; Zida et al., 2020.
TABLE 2 Infestation rate (number of fly/kg of fruit) of host plants of the Tephritidae of economic importance in Mayotte. Species names are
ordered according to their plant family.
Species name
B. dorsalis C. capitata D. ciliatus D. etiennellus N. cyanescens
2012–
2014
2019–
2021
2012–
2014
2019–
2021
2012–
2014
2019–
2021
2012–
2014
2019–
2021
2012–
2014
2019–
2021
Annona muricata _0.94_0_0_0_ 0
Citrus reticulata 00.190000000 0
Citrus sinensis 0.71 0.05 0 0 0 0 0 0 0 0
Ficus sycomorus _ 18.90 _ 0 _ 0 _ 0 _ 0
Mangifera indica 5.14 13.42 0 0 0 0 0 0 0 0
M. indica small fruits
(<50 g)
251.46 14.57 0 0 0 0 0 0 0 0
M. indica large fruits
(>50 g)
2.50 13.21 0 0 0 0 0 0 0 0
Psidium guajava 021.500000000 0
Terminalia catappa 3.21 00000000 0
Mimusops comorensis _ 0 _ 429.41 _ 0 _ 0 _ 0
Passiflora suberosa _ 0 _ 684.21 _ 0 _ 0 _ 0
Citrullus lanatus 0 _ 0 _ 29.40 _ 2.45 _ 0 _
Cucumis anguria 0 0 0 0 24.77 0 0 0 0 0
Cucumis melo 0 0 0 0 17.39 0 2.00 0 0 0
Cucumis sativus 0 0 0 0 98.76 7.67 6.64 0.49 0 0
Cucurbita moschata 0 0 0 0 135.42 214.82 17.99 0 0 0
Cucurbita pepo 0 0 0 0 261.67 11.27 0 0 0 0
Luffa acutangula 0 _ 0 _ 78.717 _ 0 _ 0 _
Sechium edule 0 _ 0 _ 31.26 _ 0 _ 0 _
Capsicum sp. 0 0 12.56 100.88 0 0 0 0 37.68 10.73
Lycopersicon
esculentum
00000000200.97 65.13
Solanum aethiopicum 0 _ 0 _ 0 _ 0 _ 391.89 _
Solanum melongena 0000000072.73 37.54
Solanum nigrum 000000002200.00 0
Note: _, No collection of this fruit was done.
6MOQUET ET AL.
values. We designed the food web for each period with the ‘bipartite’
package from a matrix of interactions among host plants and emerging
fruit fly species.
RESULTS
Tephritidae species richness
Six fruit fly species were collected during the sampling: B. dorsalis,
C. capitata, D. ciliatus, D. etiennellus, N. cyanescens and T. nigerrimum.
The latter was only observed once on an undetermined fruit on a
Solanaceae vine in the 2019–2021 sampling period.
Host range
Potential host fruit was available all year round (Appendix S1,
Figure S3). However, in some years of the study (e.g., 2019), less fruit
was available for sampling from April to August, probably because of
the rainfall deficit (Appendix S1, Figure S1).
Bactrocera dorsalis was observed on seven host plant species
(Table 2): soursoup (Annona muricata), tangerine (C. reticulata), orange
(C. sinensis), mango (M. indica), sycamore fig (Ficus sycomorus), guava
(Psidium guajava) and tropical almond (T. catappa). It is worth noting
that B. dorsalis emerged from tropical almonds during the first sam-
pling campaign, but not in 2019–2021, despite the fact that we col-
lected twice as many samples (Table 1) and took samples on five
different occasions (Appendix S1, Figure S3).
Ceratitis capitata was found in three host plant species belonging
to different families: corky passion fruit (Passiflora suberosa), Comoros
mimusops (Mimusops comorensis) and chilli pepper (Capsicum sp.).
Dacus ciliatus was observed in all cultivated Cucurbitaceae (Cucur-
bita moschata,C. pepo and C. sativus), and D. etiennellus was found in
the fruit of muskmelon (Cucumis melo), cucumber (C. sativus), squash
(C. moschata) and calabash (Lagenaria siceraria).
Neoceratitis cyanescens was observed in large numbers on culti-
vated Solanaceae, such as chilli pepper (Capsicum sp.), tomato
(L. esculentum), Ethiopian nightshade (Solanum aethiopicum) and egg-
plant (Solanum melongena).
Infestation rate
There was no significant difference (z=1.232; p=0.218) in the
infestation rate of B. dorsalis for the two sampling periods (2012–
2014 and 2019–2021, Table 2).
Infestation rates and the proportion of fruit infested by B. dorsalis
varied depending on the host plant. Total infestation rates ranged
from 0.10 fly/kg for C. sinensis to 20.84 flies/kg for P. guajava.In
2019–2020, the proportion of infested fruit ranged from 0.9% for
C. reticulata to 12.0% for P. guajava (Table 3). For M. indica,we
observed similar results in small fruit (<50 g) and in large fruit (>50 g).
The infestation rate was 11.71 flies/kg and around 5.1% of fruit was
infested.
Ceratitis capitata had a high infestation rate in two wild plant spe-
cies: Passiflora suberosa (684.2 flies/kg) and Mimusops comorensis
(429.4 flies/kg, Table 2). In 2019–2020, the proportion of infested
fruit ranged from 31.3% to 13.0% for P. suberosa and Capsicum sp.,
respectively (Table 3).
Dacus ciliatus and N. cyanescens had high infestation rates in
Cucurbitaceae (from 2.51 to 214.82 flies/kg) and Solanaceae (from
13.67 to 448.98 flies/kg), respectively (Table 2). In 2019–2020, the
proportion of fruit infested by D. ciliatus ranged from 12.5% for Cucu-
mis sativus to 50.0% for Cucurbita pepo. For N. cyanescens, the propor-
tions of infested fruit were 1.1% for Capsicum sp., 30.0% for
S. melongena and 47.58% for L. esculentum (Table 3).
Parasitoids
Only 11 individuals from one parasitoid species were observed in
samples from 2019 to 2021. We identified the species Psyttalia
insignipennis using taxonomic criteria. They emerged from six Solana-
ceae fruit (S. melongana) from two sites and probably parasitised
N. cyanescens. The parasitism rate was 0.3%.
Network
For the two periods studied, we observed three compartments in our net-
works: (i) one with B. dorsalis, (ii) one with Dacus species and (iii) one with
C. capitata and N. cyanescens, which only shared one host plant, Capsicum
sp. (Figure 2). The connectance, the number of links per species and the
cluster coefficient were lower than expected with the null model for both
networks (Table 4,p< 0.05). Nestedness and H2’were higher than
expected with the null model for the two networks (Table 4,p< 0.05).
DISCUSSION
Our study focuses on the community of Tephritidae species of socio-
economic importance between 5 and 14 years after the B. dorsalis
invasion was reported in 2007 in Mayotte (De Meyer et al., 2012).
Our main result shows that the infestation rate of polyphagous fruit
fly species, especially B. dorsalis, is unexpectedly low in the fruit sam-
ples, regardless of the year of collection.
Tephritidae richness and host range
We found six fruit fly species during this study, including four regu-
larly detected species: D. ciliatus, N. cyanescens, C. capitata and
B. dorsalis. Of the nine species identified on the island in 2012
(de Meyer et al., 2012), D. vertebratus, D. bivittatus and C. malgassa
were not observed in our samples.
FRUIT FLY INFESTATION RATE IN MAYOTTE 7
TABLE 3 Proportion of infested fruits according to fruit fly species and host plants for samples collected in 2019–2020.
Species name B. dorsalis C. capitata D. ciliatus D. etiennellus N. cyanescens
Psidium guajava 0.120 0 0 0 0
Annona muricata 0.083 0 0 0 0
Mangifera indica 0.051 0 0 0 0
M. indica small fruits (<50 g) 0.057 0 0 0 0
M. indica large fruits (>50 g) 0.044 0 0 0 0
Citrus reticulata 0.001 0 0 0 0
Ficus sycomorus 0.026 0 0 0 0
Citrus sinensis 0.011 0 0 0 0
Passiflora suberosa 0 0.313 0 0 0
Mimusops comorensis 0 0.187 0 0 0
Capsicum sp. 0 0.130 0 0 0.011
Cucurbita moschata 0 0.268 0 0
Cucurbita pepo 0 0 0.500 0 0
Cucumis sativus 0 0 0.125 0.125 0
Lycopersicon esculentum 0 0 0 0 0.476
Solanum melongena 0 0 0 0 0.300
FIGURE 2 Bipartite network analysis of host–fruit fly associations based on infestation data in Mayotte between (a) 2012 and 2014 and
(b) between 2019 and 2021. Edge width is dependent on infestation rate. We represented each network compartment with different colours.
8MOQUET ET AL.
Dacus ciliatus was observed on Cucurbitaceae. This species is
widespread in Africa (De Meyer et al., 2010) and is also present on
other Indian Ocean islands, such as Comoros, Mauritius and La
Réunion (Hassani et al., 2016; Moquet et al., 2021; Sookar
et al., 2021). It was the main species responsible for damage observed
on Cucurbitaceae in Mayotte during our surveys. For example, in
2019–2021, almost 25% of squashes were infested by D. ciliatus. This
species shared host plant species with D. etiennellus, which was much
less abundant in the host plant species studied. Dacus ciliatus remains
the fruit fly species with the greatest economic impact in Mayotte
because it causes damage to Cucurbitaceae. Control involves heavy
chemical pesticide use because there are no effective trapping sys-
tems (Alagarmalai et al., 2009; Manrakhan et al., 2017). Trials are
being conducted using mechanical methods of protection, such as
nets or screenhouses (Vanhuffel & Huat, 2019).
Neoceratitis cyanescens species was observed on Solanaceae with a
high level of infestation. Almost 50% of tomatoes were infested by this
species, which is the main threat to tomatoes in Mayotte. This oligolec-
tic species, which originates from the islands in the Indian Ocean, also
has a major impact on tomato production in Madagascar and surround-
ing islands (Hassani et al., 2022;Moquetetal.,2021; Rasolofoarivao
et al., 2021; Sookar et al., 2021). As far as D. ciliatus is concerned, other
agroecological techniques of control must be developed, for example,
sanitation and/or nets or screenhouses are being considered.
Two polyphagous species were observed: B. dorsalis and
C. capitata. The first was present in cultivated host plants, such as
M. indica and C. sinensis, while the second was particularly present in
wild species, such as M. comorensis and Passiflora suberosa. Despite
the capacity of these two species to infest similar host plants in similar
environments (Franck & Delatte, 2020), we did not observe overlap-
ping host ranges in Mayotte. Bactrocera dorsalis and C. capitata were
in a distinct compartment in the bipartite network (Figure 2). This
result could be related to the niche partitioning after the B. dorsalis
invasion or due to other unexplained factors. Interspecific competition
and climatic niche partitioning have been documented in several stud-
ies, where B. dorsalis largely displaced resident Ceratitis species (Ekesi
et al., 2009; Hassani et al., 2016; Moquet et al., 2021; Vargas
et al., 1995). This could be one of the factors that potentially plays a
role in the current host ranges observed for the two species.
The networks of host–fruit fly interactions were highly compart-
mentalised and specialised with low complexity for the two studied
periods. We observed three compartments: one with B. dorsalis, one
with C. capitata and N. cyanescens and one with the Dacus species.
Consequently, the connectance measuring the proportion of realised
interactions, among all the possible interactions in a network, was
lower than expected randomly. The H20, a network-level measure of
specialisation, was higher than expected randomly. This type of com-
partmentalised plant–fruit fly web structure was observed in a non-
pest Dacine (Tephritidae) in a New Guinea rainforest (Novotny
et al., 2005). However, this contrasts with the majority of studies on
invasive and agronomic fruit fly pests, where the presence of general-
ist species appears to induce networks with high connectance
(Charlery de la Masselière et al., 2017; Moquet et al., 2021,2023).
Infestation rate of Bactrocera dorsalis
Of the 84 plant species collected during our field sampling, 49 are
recorded in the literature as host plants for B. dorsalis. However, in
our study, only seven were infested by B. dorsalis. Observed infesta-
tion rates were low, even for fruit species recognised as preferential
hosts (>100 flies per kilogram of fruit, Appendix S2), according to Fol-
lett’s categories (Follett et al., 2021), for example: mango (11.71 flies/
kg), guava (20.84 flies/kg) and tropical almond (0.97 fly/kg). Moreover,
in 2019–2020, the proportion of infested fruit ranged from 0.9% for
tangerine to 12% for guava. For mangoes, the proportion of infested
fruit was 5.08%, with similar results on small and large fruit. Our result
suggests that the observed impact of B. dorsalis was weak compared
with other regions invaded by B. dorsalis (Appendix S2). In comparison,
other studies in the Indian Ocean showed a higher proportion of
infested fruits, with 45.5% of mangoes infested in La Réunion
(Moquet et al., 2021), 33% in the Comoros (Hassani et al., 2016) and
22% in Madagascar (Rasolofoarivao et al., 2021). Moreover, in our
study, we did not observe significant differences in infestation rates
between the first and second sampling periods. Bactrocera dorsalis
populations seemed to stabilise at low incidence during our sampling
campaigns (5 and 12 years after the B. dorsalis invasion). Many
hypotheses, which are not incompatible, could explain these results:
(i) poor climatic conditions during sampling, (ii) an equilibrium at low
incidence was reached, (iii) a long lag phase before population growth
and (iv) invasion by B. dorsalis involved a different, potentially less
competitive strain than those found on surrounding islands.
TABLE 4 Network indices calculated from bipartite networks between fruit flies and host plant species in Mayotte between 2012 and 2014
and between 2019 and 2021.
Network indices 2012–2014 p-value 2019–2021 p-value
Connectance 0.27 <0.001 0.26 <0.001
Links per species 1.04 <0.001 0.85 <0.001
Number of compartments 3.00 NA 3.00 NA
Cluster coefficient 0.20 NA 0.20 <0.001
Nestedness 37.9 <0.001 38.3 <0.001
H200.84 <0.001 0.98 <0.001
FRUIT FLY INFESTATION RATE IN MAYOTTE 9
Abiotic variables, such as weather parameters, are known to influ-
ence B. dorsalis population dynamics (Chen et al., 2006; Hassani
et al., 2022; Kamala Jayanthi & Verghese, 2011). Previous studies
showed positive correlations between B. dorsalis population size and
rainfall, as well as maximum and minimum temperatures (Kamala
Jayanthi & Verghese, 2011; Shukla & Prasad, 1985). Our data do not
allow to observe seasonal fluctuations linked to variations in tempera-
ture and humidity. Despite a less favourable cold dry season, the tem-
peratures and rainfall recorded by MétéoFrance (temperature
between 21.8 and 34C, rainfall between 0.4 and 479 mm,
Appendix S1, Figure S1) seem to correspond to suitable conditions for
the development of B. dorsalis (De Villiers et al., 2015) and, therefore,
do not explain the low infestation rates.
After the expansion stage of a biological invasion event, popula-
tion densities are supposed to reach an equilibrium, when popula-
tions of the invasive species are regulated by interspecific
interactions (competition with other fruit flies, natural enemies,
etc.),orlimitedbythecarryingcapacityofthenewenvironment
(Büyüktahtakın&Haight,2018). In our study, the first scenario is
less probable because no parasitoids have yet been deliberately
introduced into Mayotte. We only detected one indigenous parasit-
oid species in low abundance (Psyttalia insignipennis with a parasit-
ism rate of 0.3%), which is not known to infest Bactrocera species.
Moreover, we noticed the absence of one of the major parasitoid
species often used to control B. dorsalis: Fopius arisanus (Rousse
et al., 2005). The competition we observed appeared to be weak.
Only C. capitata, another polyphagous species, may compete for lar-
val resources. However, this species also had a low infestation rate
in Mayotte and we did not observe an overlap in the host range
with B. dorsalis. Thus, we cannot rule out the possibility that these
phenomena could be the result of strong competition shortly after
invasion. In La Réunion, for example, competition led to a shift in
the host range and spatial distribution of resident fruit fly species
less than 2 years after the B. dorsalis invasion. Similar observations
were noticed before in Madagascar, Comoros, Kenya or Hawaii
(Hassani et al., 2016;Keiseretal.,1974; Mwatawala et al., 2009;
Rasolofoarivao et al., 2021). Following the invasion, the host range
of C. capitata was significantly reduced and the species was only
found in host plants with small berries, which were rarely infested
by B. dorsalis (Moquetetal.,2021). Similarly, 5–12 years after the
B. dorsalis invasion in Mayotte, C. capitata was only found in fruit of
Passiflora suberosa, Mimusops comorensis and Capsicum sp.
The carrying capacity of the new environment could depend on
temporal variations in resource availability. Many studies show a rela-
tion between the fruiting period and fruit fly abundance (Abro
et al., 2021; Hassani et al., 2016; Tasnin et al., 2021; Theron
et al., 2017). Generally, despite a temporary decrease in the popula-
tion during the season when resources are scarce, the population rises
during the fruiting period of the main host plants, especially mango
(Bota et al., 2018; Motswagole et al., 2019). Tropical fruit flies have
endogenous mechanisms, such as variation in adult longevity and sea-
sonal fecundity to cope with changes in the available breeding
resources (Clarke et al., 2022; Tasnin et al., 2021). Thus, although the
decrease in the availability of fruit resources during some parts of
the year could influence temporal population dynamics, it is unlikely
to explain the overall low infestation rate observed. Indeed, little fruit
is available for B. dorsalis in Mayotte, even during the favourable
periods. We observed that farmers harvested a great deal of fruit
before ripening, either because they eat unripe fruit (e.g., banana or
mango (Weibel, 1997)), or to prevent human theft and avoid damage
by feeding vertebrates (SISE/DAAF Mayotte, 2017). When ripe, the
brown lemur (Eulemur fulvus, Nègre et al., 2006) or the flying fox (Pter-
opus seychellensis comorensis, Trewhella et al., 2001) eat fruits rapidly,
causing considerable losses to farmers (SISE/DAAF Mayotte, 2017).
However, this may be good for orchard sanitation, by limiting fruit
availability for B. dorsalis. It has already been shown that frugivorous
predators could be natural enemies of Tephritidae larvae
(Drew, 1987). Sanitation is known to be the key to effective inte-
grated pest management (IPM) to control B. dorsalis (Vargas
et al., 2016). This could partly explain the low infestation rate
observed in Mayotte.
Another hypothesis is that B. dorsalis has not yet entered the
expansion stage of its biological invasion. In some cases, demographic
processes can be more complex than the exponential growth of pests
after an introduction. There may be a time lag when the exotic species
persists in relatively low numbers before population growth
(Crooks, 2005). In Tephritidae, a lag phase lasting a number of decades
has already been observed for C. capitata in a fragmented landscape
in Kenya (Copeland et al., 2002). Many mechanisms can account for
long lags, such as intraspecific interactions (Allee effect) or genetics
(Crooks, 2005). If B. dorsalis in Mayotte is in the lag phase, rather than
the equilibrium phase, it is important to focus on monitoring popula-
tion levels to ensure a timely response in the event of a sudden popu-
lation increase. Lags in population growth and range expansion can
impact decision-making processes because the possible consequences
of the invasion are underestimated.
We considered a further hypothesis, namely, the genetic origin of
the B. dorsalis population that invaded Mayotte could be different to
that of the population in the Mascarenes (Deschepper et al., 2022).
This, combined with the impact of several bottlenecks on the invasive
population, could have resulted in the selection of less fit populations
(potentially impacted by Allee effects, Stephens & Sutherland, 1999)
than those found in other invaded countries. If this is the case, it is
important to prevent invasion by a new more virulent B. dorsalis strain
from other parts of the Indian Ocean.
CONCLUSION
In Mayotte, the infestation rate of B. dorsalis was low compared with
other regions and had less impact on cultivated species. It is essential
to understand why the situation in Mayotte differs from that in other
invaded areas. Additional studies are required to test each hypothesis
in order to explain the low abundance of B. dorsalis. This could provide
an important contribution to help manage and regulate the species,
which is expanding its geographic range.
10 MOQUET ET AL.
AUTHOR CONTRIBUTIONS
Laura Moquet: Conceptualization; data curation; formal analysis;
investigation; methodology; visualization; writing –original draft;
writing –review and editing. Tim Dupin: Investigation; methodology;
writing –review and editing. Louis Maigné: Investigation; methodol-
ogy; writing –review and editing. Joel Huat: Conceptualization; fund-
ing acquisition; project administration; writing –review and editing.
Thomas Chesneau: Investigation; methodology; writing –review and
editing. Hélène Delatte: Conceptualization; investigation; methodol-
ogy; supervision; validation; writing –review and editing.
ACKNOWLEDGEMENTS
We would like to thank Bryce Bouvard, the National Public Institution
of Mayotte and the Direction of Food, Agriculture and Forest of May-
otte (DAAF) for their contribution to sample collection and the organi-
sation of missions in Mayotte. Many thanks to all farmers, who
allowed us to sample fruit on their plots of land. Thanks to Dr. N. C.
Manoukis and the two other anonymous reviewers for their valuable
comments on the first versions of the manuscript. The authors
acknowledge the Plant Protection Platform (3P, IBISA) for its support.
FUNDING INFORMATION
This research was funded by CIRAD, the French Ministry of Agricul-
ture (MAAF), the INNOVEG project (Network for Innovation and
Transfer in Agriculture, RITA), the Région Réunion and the European
Union: European Agricultural Funds for Rural Development (EAFRD)
and European Funds for Rural Development (EFRD).
CONFLICT OF INTEREST STATEMENT
The authors declare that they have no conflict of interest in this
publication.
DATA AVAILABILITY STATEMENT
Data are available in CIRAD Dataverse (https://dataverse.cirad.fr)
https://doi.org/10.18167/DVN1/HXOZ9Z
ORCID
Laura Moquet https://orcid.org/0000-0001-7873-2218
Joel Huat https://orcid.org/0000-0002-8271-1652
Hélène Delatte https://orcid.org/0000-0001-5216-5542
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SUPPORTING INFORMATION
Additional supporting information can be found online in the Support-
ing Information section at the end of this article.
Appendix S1. Context of the study of the host range of fruit flies in
Mayotte in 2012–2014 and 2019–2021: Meteorological data
(Figure S1), land use (Figure S2) and phenology of host plants
(Figure S3).
Appendix S2. A literature review of the infestation of Mangifera indica,
Psidium guajava and Terminalia catappa by B. dorsalis (Table S1).
How to cite this article: Moquet, L., Dupin, T., Maigné, L.,
Huat, J., Chesneau, T. & Delatte, H. (2024) A study on fruit fly
host range reveals the low infestation rate of Bactrocera
dorsalis (Tephritidae) in Mayotte. Agricultural and Forest
Entomology,1–13. Available from: https://doi.org/10.1111/
afe.12614
FRUIT FLY INFESTATION RATE IN MAYOTTE 13
