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89
6AReview of the Natural
Host Plants of the
Anastrepha fraterculus
Complex in the Americas
Vicente Hernández-Ortiz*, Nancy Barradas-Juanz,
and Cecilia Díaz-Castelazo
Abstract There is now enough support for the hypothesis that nominal Anastrepha fraterculus
is a complex of cryptic species that are currently recognized, using morphometric procedures,
as eight morphotypes that probably correspond to different biological species. In addition to this
variability, there is also evidence that this nominal species presents important variation in its
range of preferential host use. The aim of this chapter is to provide a comprehensive understand-
ing of the natural host plants used by the nominal A. fraterculus under natural eld conditions.
This was accomplished through a bibliographic examination of information from the original
sources of host plants recorded for this y species. A total of 200 references from all regions of
the Americas were examined. Data useful to the analysis were captured in a database incorporat-
ing information pertaining to host identity, original source of data, and location of distribution,
where available. The list of host plants for the A. fraterculus complex comprised 177 species
belonging to 40 plant families, which together accounted for 1,622 documented reports. The
most highly represented families were Myrtaceae (27.1%), Rosaceae (11.9%), and Rutaceae
(8.5%). The Myrtaceae exhibited a high percentage (>90%) of native species in contrast to the
higher proportions of exotic species presented in the other families. Guava was the only com-
mon host shared by different populations throughout the tropical and subtropical landscapes of
the Americas. The highest number of hosts was recorded in Brazil (121), followed by Argentina
(40), Ecuador (40), Colombia (38), Venezuela (24), and Mexico (19). The landscapes occupied
by different populations of this nominal species presented some preferential patterns in terms
CONTENTS
6.1 Introduction ............................................................................................................................90
6.2 Methods and Analysis ............................................................................................................ 91
6.3 Results ..................................................................................................................................... 91
6.3.1 Diversity of Host Plants .............................................................................................. 91
6.3.2 Host Origin ............................................................................................................... 109
6.3.3 Geographical Distribution ........................................................................................ 111
6.4 Remarks and Conclusions ..................................................................................................... 113
Acknowledgments .......................................................................................................................... 114
References ...................................................................................................................................... 114
* Corresponding author.
90 Area-Wide Management of Fruit Fly Pests
of resource use. This reinforces the hypothesis of distinct taxonomic entities because most of
the plants are present throughout the range but are not found to be common hosts to all of the
y populations. In this context, the potential application of the sterile insect technique (SIT) in
certain geographical areas requires knowledge of the particular hosts consumed by the target
species.
6.1 INTRODUCTION
TheSouth American fruit y (SAFF), Anastrepha fraterculus (Wiedemann, 1830), is widely dis-
tributed throughout the Americas; from the southern United States (Texas), through Mexico, Central
America and South America to Argentina (Hernández-Ortiz& Aluja 1993). Based on previous
research under different approaches, including the use of karyotypes, molecular DNAsequences,
reproductive isolation, pheromone proles, or even integrative approaches (reviewed in Selivon
etal. 2004, 2005a, 2005b, Vera etal. 2006, Cáceres etal. 2009, Vaníčková etal. 2015, Dias etal.
2016, and others), there is nowenough support for the hypothesis that this nominal species in fact
comprises a complex of cryptic species that are currently recognized using morphometric proce-
dures as eight morphotypes, most likely corresponding to different biological species (Hernández-
Ortiz etal. 2004, 2012, 2015).
There is also evidence that some populations exhibit important differences in the host range
used, using preferential hosts at the regional scale. Moreover, their pest status can differ depending
on the geographical area in which they occur; for instance, Baker etal. (1944) showed that citrus
fruits such as oranges were unsuitable hosts for Mexican SAFF females and Aluja etal. (2003a)
later conrmed that fruits of Citrus sinensis and Citrus paradisi in Mexico are notinfested, either
in the eld or under laboratory-induced conditions. Incontrast, studies in South America revealed
that citrus fruits are common hosts for Brazilian populations of the SAFF (Malavasi etal. 1980,
Zucchi 2007, and others). Inthis regard, there are some highly questionable reports indicating that
A. fraterculus is able to infest citrus fruits in Guatemala (Eska & Cunningham 1987, Eska 1990).
Thisvariation in the range of host plants consumed in different regions of the neotropics presents a
problem in terms of enforcing effective quarantine procedures.
Historical accounts indicate that nearly 150 plant species host larvae feeding on their fruits across
the Americas (Norrbom 2004). However, many of these records were derived from observations under
laboratory conditions or copied from previous literature reports, and some fail to specify an explicit
location, or even country, or there is some uncertainty regarding the y or host plant identity. Various
studies at the regional level provide information on the population dynamics of the SAFF derived
from specimens caught in traps baited with food attractants. These traps are usually hung on fruit
trees of commercial importance; however, this does notnecessarily demonstrate that these y larvae
infest such fruits under natural conditions. Thisinformation has led to the identication of three key
problems of the phytosanitary measures that must be applied: (1) the presence of erroneous records
featuring wrongly identied or unconrmed hosts; (2) misidentication of the taxonomic species
because other wild y species are also attracted to the food baits; and (3) misinterpretation of dis-
tribution patterns based on records with inaccurate locations. As a consequence, implementation of
quarantine measures and methods of integrated pest management (IPM) based on such erroneous or
ambiguous information can lead to poor planning of control strategies based on concepts of y-free
or low prevalence areas, or even the application of the sterile insect technique (SIT) in specic geo-
graphical areas, which requires accurate identication of the hosts consumed by the target species.
Thisstudy therefore aims to provide a comprehensive understanding of the host plants used as natu-
ral food resources by A. fraterculus. Thereview focuses particularly on records produced under eld
conditions to conduct an analysis of botanical families and host species using current botanical nomen-
clature and homogenizing past and current species names, as well as examining their occurrence in dif-
ferent countries using the information available in the literature produced over nearly a century. Thisis
a critical rst step to elucidate the host range of the SAFF across several regions of the neotropics.
91AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
6.2 METHODS AND ANALYSIS
Information was sought in the original published sources of recorded host plants of the nominal
species A. fraterculus. Thesearch was largely conducted in articles published in scientic journals,
books, and unpublished dissertations available online. The main requirements of credibility for
each record were inclusion of the full identity of the host plant, as well as the occurrence of fruit
infestation under natural conditions in the eld, with no ambiguity regarding the recorded location
(to country level at least). We therefore excluded doubtful reports, those obtained under laboratory
conditions, and those in which the accurate identity of the fruit y species could notbe veried.
Nearly 170 references as primary sources of information from all regions of the Americas were
examined. All information useful for the analysis was captured in a database, including the host
identity, botanical family, original source of data, country, state or province, locality, coordinates
and collection date, where available. If a single publication reported the same host in several loca-
tions, each report was considered a separate record in the database. Furthermore, each host plant
was classied as native or exotic based on its origin because many were species that had been intro-
duced to the Americas.
Afew records were omitted from the analysis, even though they were obtained under natural
conditions. Thiswas the case of two historical records for Annona cherimola Mill. in Arica (Chile)
because A. fraterculus does not occur in that country since its eradication in 1964. Inaddition,
other reports for several countries of Rubus spp., as well as Ravenia wampi Oliv (Rutaceae) and
Cyphomandra betacea (Cav.) Stendtn. (Solanaceae) for Brazil were also excluded from the analysis
because their status as unresolved names prevented conrmation of their identity and origin.
Because some original names have changed because publication in the primary source, the
attached list presents a single identity for each host plant. Thenomenclature used for the scien-
tic names of plants was updated following the classication of “ThePlant List” (2013) to avoid
duplication or potential synonymies among species, as well as to facilitate their correct assignment
to the corresponding family. Thelist of host plants of A. fraterculus (sensu lato) is organized and
presented by host family and specic binomial name, with the authorship, origin, key code by host
species, total number of records and countries where this fruit y has been reported, as well as the
original source of information (Table6.1).
To analyze host plant trends, interaction networks depicting the associations of A. fraterculus
and its host plant species across the Americas were evaluated, considering the origin of the hosts
(native or exotic) and their plant families. Acluster analysis was also conducted based on the fre-
quency of records of hosts throughout the countries. All analyses were executed using the “plot-
web function” of the “bipartite” package (Dormann& Gruber 2009) in “R software” (RCore
Team 2014).
6.3 RESULTS
6.3.1 dIversIty oF host plants
As a result of the examination of natural host plants of A. fraterculus throughout the Americas, 177
host plants belonging to 40 plant families were recorded, which together accounted for 1,622doc-
umented reports in different localities. The plant families with highest species richness were
Myrtaceae (48), Rosaceae (20), Rutaceae (14), Leguminosae (11), Sapotaceae (10), Anacardiaceae
(8), and Annonaceae (7), which together represented nearly 66% of all recorded hosts. Forinstance,
the family Myrtaceae was represented by at least 10 different genera, having the highest species
richness. The highest frequency of records was found for Psidium guajava (298), Eugenia uni-
ora (48), Syzygium jambos (35), Acca sellowiana (31), and Psidium cattleianum (22). Myrtaceae
is therefore the most important food plant family widespread throughout the distribution range of
the SAFF.
92 Area-Wide Management of Fruit Fly Pests
TABLE 6.1
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Actinidiaceae Actinidia deliciosa
(AChev) CS Liang&
AR Ferguson
Exotic Ac-del 2 Bra Hickel and Schuck (1993)
Anacardiaceae Anacardium
occidentale L.
Native An-occ 2 Bra, Gua Eska and Cunningham (1987), Jesus-Barros etal. (2012)
Anacardiaceae Byrsonima crassifolia
(L.) Kunth
Native By-cra 1 Bra Jesus-Barros etal. (2012)
Anacardiaceae Mangifera indica L. Exotic Ma-ind 46 Arg, Bra, Col, Ecu,
FrG, Gua, Mex,
Per, Ven
Aluja etal. (1987), Alvarenga etal. (2009), Araujo (2015), Boscán de Martínez and
Godoy (1996), Campos (1960), Caraballo (1981), Eska (1990), Eska and
Cunningham (1987), Ferreira etal. (2003), Gonzalez Mendoza (1952), Guagliumi
(1966), Hernández-Ortiz etal. (2012), Korytkowski and Ojeda Peña (1968),
Korytkowski and Ojeda Peña. (1969), Malavasi etal. (1980), Marchiori etal.
(2000), Marinho (2004), Molineros etal. (1992), Núñez Bueno (1981), Ovruski
etal. (2003), Putruele etal. (1996), Raga etal. (2011), Sá etal. (2008), Silva
(1993), Silva etal. (1996), Stone (1942b), Tigrero (2009), Uramoto (2002),
Uramoto etal. (2004), Vayssières etal. (2013), Veloso etal. (2000), Zucchi (2000)
Anacardiaceae Spondias dulcis
Parkinson
Exotic Sp-dul 5 Bra Araujo (2015), Souza-Filho etal. (2000), Uramoto (2002), Uramoto etal. (2004)
Anacardiaceae Spondias mombin L. Native Sp-mom 12 Bra, Col, Mex, Per,
Ven
Aguiar-Menezes etal. (2007), Almeida (2016), Boscán de Martínez and Godoy
(1996), Gonzalez Mendoza (1952), Jesus-Barros etal. (2012), Korytkowski and
Ojeda Peña (1968), Korytkowski and Ojeda Peña (1969), Lemos (2014), Lemos
etal. (2017), McPhail and Bliss (1933), Sarmiento etal. (2012)
Anacardiaceae Spondias purpurea L. Native Sp-pur 23 Bra, Ecu, Pan, Per Aguiar-Menezes etal. (2001), Aguiar-Menezes etal. (2007), Alvarenga etal. (2000,
2009), Bressan and Da Costa Teles (1991), Campos (1960), Korytkowski and
Ojeda Peña (1968), Korytkowski and Ojeda Peña. (1969), Leal etal. (2009),
Lemos (2014), Lemos etal. (2017), Malavasi etal. (1980), Pirovani (2011),
Pirovani etal. (2010), Raga etal. (2011), Sá etal. (2008), Silva etal. (2010),
Souza-Filho etal. (2000), Stone (1942b), Tigrero (2009), Zucchi (2000)
(Continued)
93AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
(Continued)
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Anacardiaceae Spondias sp. Native Sp-sp 7Bra, Col, Ecu, Mex Aluja (1984), Molineros etal. (1992), Núñez Bueno (1981), Sá etal. (2008), Santos
(2003), Silva etal. (2010), Zucchi (2000)
Anacardiaceae Spondias tuberosa
Arruda
Native Sp-tub 2 Bra Alvarenga etal. (2009), Sá etal. (2008)
Annonaceae Annona cherimola Mill. Native An-che 21 Arg, Bra, Col, Ecu,
Per
Gonzalez Mendoza (1952), Greene (1934), Hernández-Ortiz etal. (2012),
Korytkowski and Ojeda Peña (1968), Korytkowski and Ojeda Peña. (1969),
Molineros etal. (1992), Núñez Bueno (1981), Ovruski etal. (2003), Rust (1918),
Stone (1942b), Tigrero (2009), White and Elson-Harris (1992), Wille (1941)
Annonaceae Annona crassiora
Mart.
Native An-cra 2 Bra Veloso etal. (2000), Zucchi (2000)
Annonaceae Annona muricata L. Native An-mur 4 Col Gonzalez Mendoza (1952)
Annonaceae Annona rugulosa
(Schltdl.) H. Rainer
Native An-rug 2 Bra Garcia and Norrbom (2011), Marsaro Júnior (2014)
Annonaceae Rollinia emarginata
Schltdl.
Native Ro-ema 2 Bra Souza-Filho etal. (2000), Zucchi (2000)
Annonaceae Rollinia laurifolia
Schltdl.
Native Ro-lau 2 Bra Uramoto (2007), Uramoto etal. (2008)
Annonaceae Rollinia sericea (R. E.
Fr.) R. E. Fr.
Native Ro-ser 3 Bra Raga etal. (2011), Souza-Filho etal. (2000), Zucchi (2000)
Arecaceae Butia eriospatha (Mart.
ex Drude) Becc.
Native Bu-eri 1 Bra Savaris etal. (2013)
Arecaceae Syagrus romanzofana
(Cham.) Glassman
Native Sy-rom 2 Bra Araujo (2015), Uramoto (2002)
Calophyllaceae Mammea americana L. Exotic Ma-ame 2 Col, Ecu Campos (1960), Gonzalez Mendoza (1952)
Cannabaceae Celtis iguanaea (Jacq.)
Sarg.
Native Ce-igu 1 Bra Garcia and Norrbom (2011)
Caricaceae Carica papaya L. Native Ca-pap 2 Arg, Col Gonzalez Mendoza (1952)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
94 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Celastraceae Peritassa campestris
(Cambess.) A. C. Sm.
Native Pe-cam 2 Bra Veloso etal. (2000), Zucchi (2000)
Chrysobalanaceae Chrysobalanus icaco L. Native Ch-ica 2 Bra, Col Deus etal. (2013), Gonzalez Mendoza (1952)
Clusiaceae Garcinia brasiliensis
Mart.
Native Ga-bra 2 Bra Pirovani (2011), Raga etal. (2011)
Combretaceae Terminalia catappa L. Exotic Te-cat 32 Bra, CR, Ecu,
Mex, Ven
Aguiar-Menezes etal. (2007), Aluja etal. (1987, 2000), Araujo etal. (2000), Araujo
(2012), Boscán de Martínez and Godoy (1996), Boscán de Martínez etal. (1980),
Caraballo (1981), Hernández-Ortiz and Morales-Valles (2004), Hernández-Ortiz
etal. (2012), Katiyar etal. (1995), Mascarenhas (2007), Raga etal. (2011), Silva
etal. (1996), Souza-Filho etal. (2000), Steck (1991), Steck etal. (1990), Tigrero
(2009), Zucchi (2000)
Ebenaceae Diospyros kaki L. Exotic Di-kak 12 Arg, Bra Garcia and Norrbom (2011), Malavasi etal. (1980), Marsaro Júnior (2014), Nasca
etal. (1996), Ovruski etal. (2003), Segura etal. (2004, 2006), Souza-Filho etal.
(2000), Zucchi (2000)
Euphorbiaceae Alchornea latifolia Sw. Native Al-lat 2Mex Aluja (1984), Aluja etal. (1987)
Euphorbiaceae Manihot esculenta
Crantz
Native Mn-esc 1Ven Boscán de Martínez and Godoy (1996)
Juglandaceae Juglans australis
Griseb.
Native Ju-aus 6Arg Ovruski etal. (2003, 2004)
Juglandaceae Juglans neotropica
Diels
Native Ju-neo 4 Ecu, Per Korytkowski and Ojeda Peña (1969), Molineros etal. (1992), Tigrero (2009)
Lauraceae Cryptocarya
aschersoniana Mez
Native Cr-asc 2 Bra Marinho etal. (2009), Raga etal. (2011)
Lauraceae Endlicheria paniculata
(Spreng.) J. F. Macbr.
Native En-pan 1 Bra Pirovani (2011)
Lauraceae Persea americana Mill. Native Pe-ame 4 Arg, Bra Araujo (2015), Putruele etal. (1996), Rust (1918), Uramoto (2002)
Leguminosae Andira humilis Mart. ex
Benth.
Native An-hum 2 Bra Veloso etal. (2000), Zucchi (2000)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
95AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Leguminosae Inga edulis Mart. Native In-edu 8 Bra, Col, Ecu Deus etal. (2010), Gonzalez Mendoza (1952), Lemos (2014), Lemos etal. (2017),
Malavasi etal. (1980), Tigrero (2009), Zucchi (2000)
Leguminosae Inga feuilleei DC. Native In-feu 1 Ecu Tigrero (2009)
Leguminosae Inga insignis Kunth Native In-ins 1 Ecu Tigrero (2009)
Leguminosae Inga micheliana Harms Native In-mic 1 Gua Eska and Cunningham (1987)
Leguminosae Inga paterna Harms Native In-pat 1 Gua Eska and Cunningham (1987)
Leguminosae Inga sellowiana Benth. Native In-sel 1 Bra Garcia and Norrbom (2011)
Leguminosae Inga semialata (Vell.)
C. Mart.
Native In-sem 1Arg Oroño etal. (2005)
Leguminosae Inga sp. Native In-sp 4Bra, Ecu, Ven Briceño Vergara (1975), Campos (1960), Molineros etal. (1992), Raga etal. (2011)
Leguminosae Inga spectabilis (Vahl)
Willd.
Native In-spe 2 Ecu Tigrero (2009)
Leguminosae Inga vera subsp. spuria
(Willd.) J. Leon
Native In-ver 1 Col Gonzalez Mendoza (1952)
Lythraceae Punica granatum L. Exotic Pu-gra 5 Arg, Bra, Ecu Molineros etal. (1992), Nasca etal. (1996), Putruele etal. (1996), Raga etal.
(2011), Tigrero (2009)
Malpighiaceae Malpighia emarginata
DC.
Exotic Ml-ema 4 Col, Ecu Campos (1960), Pirovani (2011), Raga etal. (2011), Sá etal. (2008)
Malpighiaceae Malpighia glabra L. Native Ml-gla 13 Bra, Col, Ecu Aguiar-Menezes etal. (2001), Araujo (2015), Gonzalez Mendoza (1952), Leal etal.
(2009), Malavasi etal. (1980), Marinho (2004), Souza-Filho etal. (2000),
Uramoto (2002), Uramoto etal. (2004), Zucchi (2000)
Malvaceae Quararibea cordata
(Bonpl.) Vischer
Native Qu-cor 3 Ecu Molineros etal. (1992), Tigrero (2009)
Malvaceae Theobroma cacao L. Native Th-cac 2Ve n Caraballo (1981), Hernández-Ortiz and Morales-Valles (2004)
Melastomataceae Bellucia
grossularioides (L.)
Triana
Native Be-gro 1 FrGuy Vayssières etal. (2013)
96 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Melastomataceae Mouriri acutiora
Naudin
Native Mo-acu 1 Bra Jesus-Barros etal. (2012)
Melastomataceae Mouriri glazioviana
Cogn.
Native Mo-gla 2 Bra Uramoto (2007), Uramoto etal. (2008)
Moraceae Ficus carica L. Exotic Fi-car 16 Arg, Bra, Ecu Garcia and Norrbom (2011), Molineros etal. (1992), Nasca etal. (1996), Ovruski
etal. (2003), Putruele etal. (1996), Rust (1918), Tigrero (2009)
Moraceae Helicostylis sp. Native He-sp 3 Bra Bondar (1950), Kovaleski etal. (1999), Zucchi (2000)
Moraceae Helicostylis tomentosa
(Poepp.& Endl.)
J.F.Macbr.
Native He-tom 1 Bra Bondar (1950)
Myrtaceae Acca sellowiana
(O.Berg) Burret
Native Ac-sel 31 Arg, Bra, Col, Ecu Alberti etal. (2002), Cruz etal. (2017), Custódio etal. (2017), Garcia and Norrbom
(2011), Hernández-Ortiz etal. (2015), Hickel and Ducroquet (1994), Kovaleski
etal. (1999, 2000), Marsaro Júnior (2014), Molineros etal. (1992), Nasca etal.
(1996), Nunes etal. (2012), Ovruski etal. (2008), Pereira-Rêgo etal. (2011), Rust
(1918), Salles (1995), Schliserman etal. (2010), Segura etal. (2004, 2006),
Tigrero (2009), Yepes and Vélez (1989), Zucchi (2000)
Myrtaceae Calycolpus moritzianus
(O. Berg) Burret
Native Ca-mor 3Ven Hernández-Ortiz and Morales-Valles (2004), Hernández-Ortiz etal. (2012), Katiyar
etal. (2000)
Myrtaceae Campomanesia
adamantium
(Cambess.) O. Berg
Native Cm-ada 8 Bra Malavasi etal. (1980), Selivon (2000), Souza-Filho etal. (2000), Veloso etal.
(2000), Zucchi (2000)
Myrtaceae Campomanesia
espiritosantensis
Landrum
Native Cm-esp 2 Bra Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Campomanesia
guaviroba (DC.)
Kiaersk.
Native Cm-gua 1 Bra Zucchi (2000)
(Continued)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
97AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
(Continued)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Myrtaceae Campomanesia
guazumifolia
(Cambess.) O. Berg
Native Cm-guz 2 Bra Garcia and Norrbom (2011), Marsaro Júnior (2014)
Myrtaceae Campomanesia
lineatifolia Ruiz&
Pav.
Native Cm-lin 2 Bra Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Campomanesia
pubescens (Mart. ex
DC.) O. Berg
Native Cm-pub 2 Bra Guimarães etal. (1999), Raga etal. (2005)
Myrtaceae Campomanesia
xanthocarpa (Mart.)
O. Berg
Native Cm-xan 8 Arg, Bra Garcia and Norrbom (2011), Kovaleski etal. (1999, 2000), Malavasi etal. (1980),
Marsaro Júnior (2014), Salles (1995), Schliserman etal. (2010), Zucchi (2000)
Myrtaceae Eugenia brasiliensis
Lam.
Native Eu-bra 8 Bra Aguiar-Menezes etal. (2007), Raga etal. (2005), Steck and Malavasi (1988), Steck
etal. (1990), Stone (1942b), Uramoto (2007), Uramoto etal. (2008), Wille (1941)
Myrtaceae Eugenia dodonaeifolia
Cambess.
Native Eu-dod 1 Bra Zucchi (2000)
Myrtaceae Eugenia dysenterica
DC.
Native Eu-dys 3 Bra Silva etal. (2010), Veloso etal. (2000), Zucchi (2000)
Myrtaceae Eugenia orida DC. Native Eu-o 1Pan Stone (1942a)
Myrtaceae Eugenia gemmiora
O.Berg
Native Eu-gem 2 Bra Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Eugenia involucrata
DC.
Native Eu-inv 15 Bra Araujo (2015), Garcia and Norrbom (2011), Haji and Da Gama Miranda (2000),
Kovaleski etal. (1999, 2000), Marsaro Júnior (2014), Nunes etal. (2012), Raga
etal. (2005), Salles (1995), Uramoto (2002, 2007), Uramoto etal. (2004, 2008),
Zucchi (2000)
Myrtaceae Eugenia lambertiana
DC.
Native Eu-lam 2 Bra Raga etal. (2005), Zucchi (2000)
Myrtaceae Eugenia leitonii D.
Legrand
Native Eu-lei 2 Bra Raga etal. (2005), Zucchi (2000)
98 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Myrtaceae Eugenia myrcianthes
Nied.
Native Eu-myr 2Arg Putruele etal. (1996), Segura etal. (2006)
Myrtaceae Eugenia platyphylla
O.Berg
Native Eu-pla 3 Bra Uramoto (2007)
Myrtaceae Eugenia platysema
O.Berg
Native Eu-plt 1 Bra Uramoto etal. (2008)
Myrtaceae Eugenia punicifolia
(Kunth) DC.
Native Eu-pun 2Ven Fernandez Yepez (1953), Guagliumi (1966)
Myrtaceae Eugenia pyriformis
Cambess.
Native Eu-pyr 19 Bra Araujo (2015), Garcia and Norrbom (2011), Guimarães etal. (1999), Kovaleski
etal. (1999, 2000), Malavasi etal. (1980), Marinho (2004), Marsaro Júnior
(2014), Nunes etal. (2012), Perre (2016), Raga etal. (2005), Souza-Filho etal.
(2000), Uramoto (2002), Uramoto etal. (2004), Zucchi (2000)
Myrtaceae Eugenia stipitata
McVaugh
Native Eu-sti 8 Bra, Ecu Lemos (2014), Lemos etal. (2017), Molineros etal. (1992), Silva etal. (2010),
Tigrero (2009), Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Eugenia uniora L. Native Eu-uni 48 Arg, Bra, Gua,
Mex, Pan
Aguiar-Menezes and Menezes (2000), Aguiar-Menezes etal. (2003), Aguiar-
Menezes etal. (2007), Aluja etal. (1987), Alvarenga etal. (2009), Alvarenga etal.
(2000), Araujo (2015), Eska (1990), Garcia and Norrbom (2011), Guimarães
etal. (1999), Hernández-Ortiz etal. (2012), Malavasi etal. (1980), Marchiori etal.
(2000), Marinho (2004), Marsaro Júnior (2014), Nunes etal. (2012), Ovruski etal.
(2003, 2004), Ovruski etal. (2008), Pirovani (2011), Pirovani etal. (2010), Raga
etal. (2005), Salles (1995), Silva etal. (2010, 2011), Souza-Filho etal. (2000),
Stone (1942b), Uramoto (2002), Uramoto etal. (2004), Veloso etal. (2000),
Zucchi (2000)
Myrtaceae Myrceugenia euosma
(O. Berg) D. Legrand
Native Me-euo 3 Bra Kovaleski etal. (1999, 2000), Zucchi (2000)
Myrtaceae Myrcia clausseniana
Berg
Native Mc-cla 2 Bra Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Myrcia popayanensis
Hieron.
Native Mc-pop 1 Col Olarte Espinosa (1980)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
99AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Myrtaceae Myrcianthes fragrans
(Sw.) McVaugh
Exotic My-fra 1 Col Gonzalez Mendoza (1952)
Myrtaceae Myrcianthes pungens
(O. Berg) D. Legrand
Native My-pun 11 Arg, Bra Garcia and Norrbom (2011), Marsaro Júnior (2014), Ovruski etal. (2003, 2004,
2008), Putruele etal. (1996)
Myrtaceae Myrciaria dubia
(Kunth) McVaugh
Native Mr-dub 1 Bra Custódio etal. (2017)
Myrtaceae Myrciaria oribunda
(H.West ex Willd.)
O.Berg
Native Mr-o 1Mex Aluja etal. (2000)
Myrtaceae Myrciaria glazioviana
(Kiaersk.) G. M.
Barroso ex Sobral
Native Mr-gla 1 Bra Pirovani (2011)
Myrtaceae Myrciaria glomerata
O.Berg
Native Mr-glo 5 Bra Araujo (2015), Marinho (2004), Raga etal. (2005), Uramoto (2002), Uramoto etal.
(2004)
Myrtaceae Myrciaria strigipes
O.Berg
Native Mr-str 2 Bra Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Plinia cauliora
(Mart.) Kausel
Native Pl-cau 15 Bra Aguiar-Menezes and Menezes (2000), Aguiar-Menezes etal. (2007), Araujo (2015),
Garcia and Norrbom (2011), Marinho (2004), Pirovani (2011), Pirovani etal.
(2010), Raga etal. (2005), Salles (1995), Silva etal. (2011), Souza-Filho etal.
(2000), Uramoto (2002), Uramoto etal. (2004), Zucchi (2000)
Myrtaceae Plinia edulis (Vell.)
Sobral
Native Pl-edu 2 Bra Raga etal. (2005), Zucchi (2000)
Myrtaceae Psidium acutangulum
Mart. ex DC.
Native Ps-acu 1 Col Hernández-Ortiz etal. (2015)
Myrtaceae Psidium cattleianum
Afzel. ex Sabine
Native Ps-cat 22 Arg, Bra, Gua Aguiar-Menezes etal. (2001), Eska and Cunningham (1987), Garcia and Norrbom
(2011), Kovaleski etal. (1999, 2000), Leal etal. (2009), Manni etal. (2015),
Marinho (2004), Marsaro Júnior (2014), Nunes etal. (2012), Pereira-Rêgo etal.
(2011), Raga etal. (2005), Rust (1918), Zucchi (2000)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
100 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Myrtaceae Psidium guajava L. Native Ps-gua 298 Arg, Bol, Bra, Col,
CR, Ecu, FrG,
Gua, Mex, Pan,
Per, Ven
Adaime etal. (2017), Aguiar-Menezes and Menezes (1997, 2000), Aguiar-Menezes
etal. (2001, 2007), Alberti etal. (2002), Aluja (1984), Aluja etal. (1987, 2000),
Alvarenga etal. (2000. 2009), Araujo etal. (2000), Araujo (2012, 2015), Baker etal.
(1944), Barros (2008), Bomm etal. (2007), Boscán de Martínez and Godoy (1996),
Campos (1960), Caraballo (1981), Castañeda etal. (2010), Corsato (2004), Cruz etal.
(2017), Deus etal. (2010), Eska (1990), Eska and Cunningham (1987), Fernandez
Yepez (1953), Fischer (1934), Garcia and Norrbom (2011), Gonzalez Mendoza
(1952), Guagliumi (1966), Haji and Da Gama Miranda (2000), Hedström (1987),
Hernández-Ortiz and Morales-Valles (2004), Hernández-Ortiz and Pérez-Alonso
(1993), Hernández-Ortiz etal. (1994, 2004, 2012, 2015), Isaac (1905), Jesus-Barros
etal. (2012), Jirón and Hedström (1988), Katiyar etal. (2000), Korytkowski etal.
(2001), Korytkowski and Ojeda Peña (1968, 1969),Leal etal. (2009), Lemos (2014),
Lemos etal. (2017), Malavasi etal. (1980), Manni etal. (2015), Marchiori etal.
(2000), Marsaro Júnior (2014), Mascarenhas (2007), McPhail and Bliss (1933),
Molineros etal. (1992), Nunes etal. (2012), Núñez Bueno (1981), Nuñez Bueno
etal. (2004), Ovruski (1995), Ovruski etal. (2003, 2004, 2005, 2008, 2009),
Pereira-Rêgo etal. (2011), Perre (2016), Pirovani (2011), Pirovani etal. (2010),
Putruele etal. (1996), Querino etal. (2014), Raga etal. (2005), Sá etal. (2008), Salles
(1995), Sarmiento etal. (2012), Schliserman etal. (2010), Segura etal. (2006),
Selivon (2000), Silva etal. (2010, 2011a, 2011b), Silva and Silva (2007), Souza-Filho
(2005, 2006), Souza-Filho etal. (2009), Steck (1991), Steck etal. (1990), Stone
(1942b), Taira (2012), Taira etal. (2013), Tigrero (1998, 2009), Uchôa-Fernandes and
Zucchi (2000), Uchôa-Fernandes etal. (2002), Uramoto (2002), Uramoto etal.
(2004), Vayssières etal. (2013), Veloso etal. (2000), Wille (1937), Zucchi (2000)
Myrtaceae Psidium guineense Sw. Native Ps-gui 17 Bra, Col, Mex, Ven Aluja etal. (2000), Araujo etal. (2000), Bomm etal. (2007), Castañeda etal. (2010),
Fernandez Yepez (1953), Gonzalez Mendoza (1952), Guagliumi (1966), Hernández-
Ortiz etal. (2004), Katiyar etal. (2000), Pirovani (2011), Pirovani etal. (2010), Silva
etal. (2010, 2011), Uramoto etal. (2008), Veloso etal. (2000), Zucchi (2000)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
101AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Myrtaceae Psidium guyanense
Pers.
Native Ps-guy 1 Bra Uramoto (2007)
Myrtaceae Psidium kennedyanum
Morong
Native Ps-ken 1 Bra Uchôa-Fernandes and Nicácio (2010)
Myrtaceae Psidium myrtoides O.
Berg
Native Ps-myr 2 Bra Uramoto (2007), Uramoto etal. (2008)
Myrtaceae Psidium sartorianum
(O. Berg) Nied.
Native Ps-sar 4Mex Aluja etal. (2000), Aluja etal. (2003b)
Myrtaceae Psidium sp. Native Ps-sp 8 Arg, Bra, Ecu Molineros etal. (1992), Rust (1918), Salles (1995), Tigrero (2009), Uchôa-
Fernandes and Zucchi (2000), Uchôa-Fernandes etal. (2002)
Myrtaceae Syzygium aqueum
(Burm. f.) Alston
Exotic Sy-aqu 1 Bra Zucchi (2000)
Myrtaceae Syzygium jambos (L.)
Alston
Exotic Sy-jam 35 Bra, Ecu, Mex,
Per, Ven
Aluja (1984), Aluja etal. (1987, 2000), Araujo (2015), Baker etal. (1944), Baker
(1945), Bomm etal. (2007), Bush (1962), Campos (1960), Hernández-Ortiz etal.
(2004), Korytkowski and Ojeda Peña (1968, 1969), Marinho (2004), Molineros
etal. (1992), Perre (2016), Raga etal. (2005), Souza-Filho etal. (2000), Steck
(1991), Steck etal. (1990), Tigrero (2009), Uchôa-Fernandes and Nicácio (2010),
Uramoto (2002), Uramoto etal. (2004)
Myrtaceae Syzygium malaccense
(L.) Merr.& L. M.
Perry
Exotic Sy-mal 4 Bra, Pan Silva etal. (2010, 2011), Stone (1942b)
Oleaceae Olea europaea L. Exotic Ol-eur 1Arg Nasca etal. (1996)
Oxalidaceae Averrhoa carambola L. Exotic Av-car 18 Bra Aguiar-Menezes etal. (2001, 2007), Araujo (2015), Feitosa etal. (2007), Garcia
and Norrbom (2011), Leal etal. (2009), Malavasi etal. (1980), Marchiori etal.
(2000), Raga etal. (2011), Silva etal. (2010), Souza-Filho etal. (2000), Uramoto
(2002), Uramoto etal. (2004), Veloso etal. (2000), Zucchi (2000)
Passioraceae Passiora alata Curtis Native Pa-ala 3 Bra Pirovani (2011), Pirovani etal. (2010), Souza-Filho etal. (2000)
Passioraceae Passiora caerulea L. Native Pa-cae 3Arg Ovruski etal. (2003), Putruele etal. (1996)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
102 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Passioraceae Passiora edulis Sims Native Pa-edu 4 Bra Araujo (2015), Souza-Filho etal. (2000), Uramoto (2002), Uramoto etal. (2004)
Passioraceae Passiora sp. Native Pa-sp 3 Bra Kovaleski etal. (1999, 2000), Zucchi (2000)
Passioraceae Passiora tripartita
(Juss.) Poir.
Native Pa-tri 1 Col Castro (2009)
Picramniaceae Picramnia sp. Native Pi-sp 1 Bra Raga etal. (2011)
Rhamnaceae Ziziphus joazeiro Mart. Native Zi-joa 1 Bra Sá etal. (2008)
Rhamnaceae Ziziphus jujuba Mill. Native Zi-juj 1 Col Gonzalez Mendoza (1952)
Rosaceae Cydonia oblonga Mill. Exotic Cy-obl 6Arg, Per, Ven Guagliumi (1966), Korytkowski and Ojeda Peña (1968, 1969), Nasca etal. (1996),
Ovruski etal. (2010), Putruele etal. (1996)
Rosaceae Eriobotrya japonica
(Thunb.) Lindl.
Exotic Er-jap 75 Arg, Bra, Col, Ecu,
Mex, Per, Ven
Aguiar-Menezes and Menezes (2000), Aguiar-Menezes etal. (2007), Alberti etal.
(2002), Aluja (1984), Araujo (2015), Briceño Vergara (1975, 1979), Caraballo
(1981), Cruz etal. (2017), Fernandez Yepez (1953), Garcia and Norrbom (2011),
Gonzalez Mendoza (1952), Guagliumi (1966), Hernández-Ortiz and Morales-
Valles (2004), Hernández-Ortiz etal. (2012), Katiyar etal. (1995), Kovaleski etal.
(1999, 2000), Malavasi etal. (1980), Marinho (2004), Mascarenhas (2007),
Molineros etal. (1992), Nunes etal. (2012), Ovruski etal. (2003), Perre (2016),
Pirovani (2011), Pirovani etal. (2010), Putruele etal. (1996), Raga etal. (2011),
Salles (1995), Schliserman etal. (2010), Segura etal. (2006), Souza-Filho (2006),
Souza-Filho etal. (2000, 2009), Steck (1991), Stone (1942b), Tigrero (2009),
Uramoto (2002), Uramoto etal. (2004), Yepes and Vélez (1989), Zucchi (2000)
Rosaceae Fragaria×ananassa
(Duchesne ex Weston)
Duchesne ex Rozier
Exotic Fg-ana 1 Bra Raga etal. (2011)
Rosaceae Fragaria vesca L. Exotic Fg-ves 2 Bra Salles (1995), Zucchi (2000)
Rosaceae Malus domestica
Borkh.
Exotic Ma-dom 16 Bra, Col, Ecu, Ven Ballou (1945), Boscán de Martínez etal. (1980), Campos (1960), Caraballo (1981),
Gonzalez Mendoza (1952), Guagliumi (1966), Kovaleski etal. (1999, 2000),
Malavasi etal. (1980), Molineros etal. (1992), Perre (2016), Raga etal. (2011),
Salles (1995), Tigrero (2009), Zucchi (2000)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
103AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Rosaceae Prunus armeniaca L. Exotic Pr-arm 6Arg Nasca etal. (1996), Ovruski etal. (2003, 2004), Putruele etal. (1996), Rust (1918)
Rosaceae Prunus avium (L.) L. Exotic Pr-avi 1 Bra Garcia and Norrbom (2011)
Rosaceae Prunus domestica L. Exotic Pr-dom 17 Arg, Bra, Col, Ecu Garcia and Norrbom (2011), Guimarães etal. (1999), Kovaleski etal. (1999, 2000),
Marin Patiño (2002), Molineros etal. (1992), Nasca etal. (1996), Ovruski etal.
(2003, 2004, 2010), Raga etal. (2011), Salles (1995), Tigrero (2009), Zucchi
(2000)
Rosaceae Prunus domestica
subsp. insititia (L.)
Bonnier& Layens
Exotic Pr-dom 2Arg Putruele etal. (1996), Segura etal. (2006)
Rosaceae Prunus dulcis (Mill.)
D. A.Webb
Exotic Pr-dul 1Arg Nasca etal. (1996)
Rosaceae Prunus mume (Siebold)
Siebold& Zucc.
Exotic Pr-mum 2 Bra Perre (2016), Raga etal. (2011)
Rosaceae Prunus persica (L.)
Batsch
Exotic Pr-per 177 Arg, Bol, Bra, Col,
Ecu, Gua, Mex,
Per, Ven
Aguiar-Menezes and Menezes (1997), Alberti etal. (2002), Aluja etal. (2000),
Araujo (2015), Baker etal. (1944), Boscán de Martínez and Godoy (1996),
Boscán de Martínez etal. (1980), Briceño Vergara (1975, 1979), Campos (1960),
Caraballo (1981), Cruz etal. (2017), Eska and Cunningham (1987), Garcia and
Norrbom (2011), Gonzalez Mendoza (1952), Greene (1934), Guagliumi (1966),
Hernández-Ortiz and Morales-Valles (2004), Kovaleski etal. (1999, 2000),
Malavasi etal. (1980), Marsaro Júnior (2014), Molineros etal. (1992), Nasca etal.
(1996), Nunes etal. (2012), Ovruski (1995), Ovruski etal. (2003, 2004, 2009,
2010), Perre (2016), Putruele etal. (1996), Raga etal. (2011), Salles (1995),
Schliserman etal. (2010), Segura etal. (2004, 2006), Shaw (1947), Souza-Filho
(2006), Souza-Filho etal. (2000, 2009), Steck (1991), Tigrero (2009), Uramoto
(2002), Uramoto etal. (2004), Zucchi (2000)
Rosaceae Prunus salicina Lindl. Exotic Pr-sal 7 Arg, Bra Alberti etal. (2002), Marinho (2004), Marinho etal. (2009), Perre (2016)
Rosaceae Prunus serotina Ehrh.
ssp. capuli (Cav.)
McVaugh
Native Pr-ser 1 Gua Eska and Cunningham (1987)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
104 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Rosaceae Prunus sp. Exotic Pr-sp 5 Bra, Col Gonzalez Mendoza (1952), Malavasi etal. (1980), Salles (1995), Zucchi (2000)
Rosaceae Pyrus communis L. Exotic Py-com 18 Arg, Bra, Ecu, Per Aguiar-Menezes and Menezes (2000), Campos (1960), Garcia and Norrbom
(2011), Korytkowski and Ojeda Peña (1968, 1969), Kovaleski etal. (2000),
Molineros etal. (1992), Nasca etal. (1996), Nunes etal. (2015), Raga etal.
(2011), Salles (1995), Souza-Filho etal. (2000), Stone (1942b), Tigrero (2009),
Zucchi (2000)
Rosaceae Rubus eriocarpus
Liebm.
Native Ru-eri 3Ecu, Ven Hernández-Ortiz and Morales-Valles (2004), Tigrero (2009)
Rosaceae Rubus idaeus L. Exotic Ru-ida 1Arg Funes etal. (2017)
Rosaceae Rubus sp. ? Ru-sp 26 Arg, Bra, Col, Ecu,
Ven
Araujo (2015), Briceño Vergara (1975, 1979), Castañeda etal. (2010), Funes etal.
(2017), Hernández-Ortiz etal. (2012), Katiyar etal. (1995), Kovaleski etal. (2000),
Manni etal. (2015), Molineros etal. (1992), Núñez Bueno (1981), Salles (1995), Steck
(1991), Steck etal. (1990), Uramoto (2002), Uramoto etal. (2004), Zucchi (2000)
Rosaceae Rubus ulmifolius Schott Exotic Ru-ulm 1 Bra Raga etal. (2011)
Rubiaceae Coffea arabica L. Exotic Co-ara 51 Bra, Col, Mex, Ven Aluja (1984), Aluja etal. (1987), Araujo (2015), Boscán de Martínez etal. (1980),
Caraballo (1981), Castañeda etal. (2010), Cruz etal. (2017), Gonzalez Mendoza
(1952), Hernández-Ortiz and Morales-Valles (2004), Hernández-Ortiz etal.
(2012), Hernández-Ortiz etal. (2015), Katiyar etal. (1995), Malavasi etal. (1980),
Núñez Bueno (1981), Nuñez Bueno etal. (2004), Sarmiento etal. (2012), Souza
etal. (2005), Steck (1991), Steck etal. (1990), Stone (1942b), Torres (2004),
Uramoto (2002), Uramoto etal. (2004), Yepes and Vélez (1989), Zucchi (2000)
Rubiaceae Coffea liberica Hiern Exotic Co-lib 1Ve n Guagliumi (1966)
Rutaceae Citrus aurantiifolia
(Christm.) Swingle
Exotic Ct-arn 4 Bra Salles (1995), Zucchi (2000)
Rutaceae Citrus japonica Thunb. Exotic Ct-jap 5 Arg, Bra, Col Gonzalez Mendoza (1952), Raga etal. (2004), Rust (1918), Salles (1995), Zucchi
(2000)
Rutaceae Citrus limon (L.)
Osbeck
Exotic Ct-lim 3 Bra Salles (1995). Souza-Filho etal. (2000), Zucchi (2000)
(Continued)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
105AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Rutaceae Citrus maxima (Burm.)
Merr.
Exotic Ct-max 8 Arg, Bra, Ecu, Per Korytkowski and Ojeda Peña (1968, 1969), Malavasi etal. (1980), Raga etal.
(2004), Rust (1918), Souza-Filho etal. (2000), Tigrero (2009), Zucchi (2000)
Rutaceae Citrus medica L. Exotic Ct-med 2 Bra, Gua Eska and Cunningham (1987), Raga etal. (2004)
Rutaceae Citrus nobilis Lour. Exotic Ct-nob 1Arg Rust (1918)
Rutaceae Citrus paradisi Macfad. Exotic Ct-par 32 Arg, Bra, Ecu,
Gua, Ven
Caraballo (1981), Eska and Cunningham (1987), Molineros etal. (1992), Nasca
etal. (1996), Ovruski etal. (2003), Putruele etal. (1996), Schliserman etal.
(2010), Segura etal. (2006), Zucchi (2000)
Rutaceae Citrus reticulata
Blanco
Exotic Ct-ret 30 Arg, Bra, Col, Ecu Araujo (2015), Castañeda etal. (2010), Garcia and Norrbom (2011), Molineros
etal. (1992), Ovruski etal. (2003), Putruele etal. (1996), Raga etal. (2004),
Schliserman etal. (2010), Segura etal. (2004, 2006), Souza-Filho etal. (2000),
Tigrero (2009), Uramoto (2002), Uramoto etal. (2004)
Rutaceae Citrus sinensis (L.)
Osbeck
Exotic Ct-sin 44 Arg, Bra, Ecu,
Gua, Per
Aguiar-Menezes and Menezes (2000), Ballou (1945), Boscán de Martínez etal.
(1980), Eska (1990), Eska and Cunningham (1987), Guagliumi (1966),
Guimarães etal. (1999), Korytkowski and Ojeda Peña (1968, 1969), Malavasi
etal. (1980), Molineros etal. (1992), Ovruski etal. (2003), Pirovani (2011),
Pirovani etal. (2010), Putruele etal. (1996), Raga etal. (2004), Salles (1995),
Selivon (2000), Silva etal. (2006), Stone (1942b), Tigrero (2009), Zucchi (2000)
Rutaceae Citrus sinensis x C
reticulata
Exotic Ct-sxr 2 Bra Silva etal. (2006)
Rutaceae Citrus sp. Exotic Ct-sp 34 Bra, Col, Ecu, Ven Araujo (2012, 2015), Briceño Vergara (1979), Caraballo (1981), Gonzalez Mendoza
(1952), Molineros etal. (1992), Núñez Bueno (1981), Raga etal. (2004), Salles
(1995), Uramoto (2002), Uramoto etal. (2004), Zucchi (2000)
Rutaceae Citrus trifoliata L. Exotic Ct-tri 2 Ecu Molineros etal. (1992), Tigrero (2009)
Rutaceae Citrus x aurantium L. Exotic Ct-aur 39 Arg, Bra, Col, Ecu,
Mex, Ven
Briceño Vergara (1975), Campos (1960), Gonzalez Mendoza (1952), Hernández-
Ortiz etal. (1994, 2006), Malavasi etal. (1980), Marchiori etal. (2000), Ovruski
etal. (2003), Putruele etal. (1996), Raga etal. (2004), Schliserman and Ovruski
(2004), Schliserman etal. (2010), Segura etal. (2006), Tigrero (1998, 2009),
Yepes and Vélez (1989), Zucchi (2000)
106 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Rutaceae Ravenia wampi Oliv. ? unres.
name
1 Bra Aguiar-Menezes etal. (2001)
Salicaceae Dovyalis abyssinica
(A.Rich.) Warb.
Exotic Do-aby 2 Ecu Molineros etal. (1992), Tigrero (2009)
Salicaceae Dovyalis hebecarpa
(Gardner) Warb.
Exotic Do-heb 1Pan Stone (1942b)
Salicaceae Zuelania guidonia
(Sw.) Britton&
Millsp.
Native Zu-gui 1Mex García-Ramírez etal. (2010)
Sapindaceae Diatenopteryx
sorbifolia Radlk.
Native Di-sor 2 Bra Salles (1995), Zucchi (2000)
Sapindaceae Melicoccus bijugatus
Jacq.
Native Me-bij 1 Bra Gonzalez Mendoza (1952)
Sapindaceae Melicoccus oliviformis
Kunth
Native Me-oli 1Mex García-Ramírez etal. (2010)
Sapotaceae Chrysophyllum cainito L. Native Cs-cai 3Ecu, Ven Campos (1960), Hernández-Ortiz and Morales-Valles (2004)
Sapotaceae Chrysophyllum
gonocarpum (Mart.&
Eichler ex Miq.) Engl.
Native Cs-gon 3 Arg, Bra Oroño etal. (2005), Salles (1995), Zucchi (2000)
Sapotaceae Chrysophyllum
mexicanum Brandegee
Native Cs-mex 1 Bra Souza-Filho etal. (2000)
Sapotaceae Manilkara zapota (L.)
P. Royen
Native Ma-zap 8 Bra, Col, Ecu, Pan Araujo (2015), Campos (1960), Gonzalez Mendoza (1952), Núñez Bueno (1981),
Raga etal. (2011), Stone (1942b), Uramoto (2002), Uramoto etal. (2004)
Sapotaceae Planchonella obovata
(R. Br.) Pierre
Native Pl-obo 1 Per Korytkowski and Ojeda Peña. (1969)
Sapotaceae Pouteria caimito
(Ruiz& Pav.) Radlk.
Native Po-cai 10 Bra, Ecu Aguiar-Menezes etal. (2007), Araujo (2015), Raga etal. (2011), Souza-Filho etal.
(2000), Tigrero (2009), Uramoto (2002), Uramoto etal. (2004), Veloso etal.
(2000), Zucchi (2000)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
107AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Sapotaceae Pouteria gardneriana
(A. DC.) Radlk.
Native Po-gar 3 Bra Veloso etal. (1996, 2000), Zucchi (2000)
Sapotaceae Pouteria ramiora
(Mart.) Radlk.
Native Po-ram 3 Bra Veloso etal. (1996, 2000), Zucchi (2000)
Sapotaceae Pouteria torta (Mart.)
Radlk.
Native Po-tor 1 Bra Taira (2012)
Sapotaceae Sideroxylon capiri
(A.DC.) Pittier
Native Si-cap 1Mex Aluja etal. (1987)
Simaroubaceae Simaba guianensis
Aubl.
Native Sm-gui 1 Bra Deus etal. (2013)
Siparunaceae Siparuna guianensis
Aubl.
Native Sp-gui 1 Bra Pirovani (2011)
Solanaceae Capsicum annuum L. Native Ca-ann 1 Col Castañeda etal. (2010)
Solanaceae Cyphomandra betacea
(Cav.) Sendtn.
?unres.
name
1 Col Gonzalez Mendoza (1952)
Solanaceae Solanum
decompositiorum
Sendtn.
Native So-dec 1 Bra Pirovani (2011)
Solanaceae Solanum nudum Dunal Native So-nud 1 Col Castañeda etal. (2010)
Solanaceae Solanum quitoense
Lam.
Native So-qui 1 Col Núñez Bueno (1981)
Solanaceae Solanum triste Jacq. Native So-tri 1 Col Gonzalez Mendoza (1952)
Staphyleaceae Turpinia occidentalis
(Sw.) G. Don
Native Tu-occ 1Pan Stone (1942b)
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
(Continued)
108 Area-Wide Management of Fruit Fly Pests
Host Family Valid Scientic Name
Host
Origin
Host-
Code
Total
records Countries References
Ulmaceae Ampelocera hottlei
(Standl.) Standl.
Native Am-hot 1Mex Aluja etal. (2003b)
Urticaceae Pourouma sp. Native Po-sp 1 Bra Bondar (1950)
Vitaceae Vitis vinifera L. Exotic Vi-vin 3 Arg, Col Gonzalez Mendoza (1952), Nasca etal. (1996), Stone (1942b)
Note: Nomenclatures of scientic names are currently valid according to ThePlant List (2013).
Abbreviations: Arg, Argentina; Bra, Brazil; Bol, Bolivia; Col, Colombia; CR, Costa Rica; Ecu, Ecuador; FrG, French Guyana; Gua, Guatemala; Mex, Mexico; Pan, Panama; Per, Peru; Ven,
Venezuela.
TABLE 6.1 (Continued)
List of Host Plants for the Nominal Species Anastrepha fraterculus Reported in the Literature as Food Plants Infested in Nature
109AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
Inthe family Rosaceae, 20 species were found, with most reports featuring Prunus spp., Rubus
spp., and Eriobotrya japonica. Thefamily Rutaceae included 14 host species, almost all of which
were Citrus spp. Inthe case of the family Combretaceae, all reports were of a single host, the tropi-
cal almond Terminalia cattapa. Alarge proportion of at least 18 plant families presented only one
tothree hosts and a similarly low number of reports; thus, these could be considered rare hosts.
These included Actinidia deliciosa (Actinidaceae), Celtis iguanaea (Cannabaceae), Endlicheria
paniculata (Lauraceae), Butia eriospatha (Arecaceae), and Simaba guianensis (Simaroubaceae),
among others (Figure6.1 and Table6.1).
6.3.2 Host origin
Examination of the origin of hosts revealed that the SAFF feeds on 50 exotic plant species in
the Americas, representing 28.3% of the total number of their host plant species. Five of these,
all belonging to different families, accounted for the highest number of records: Prunus persica
(Rosaceae), Eriobotrya japonica (Myrtaceae), Coffea arabica (Rubiaceae), Mangifera indica
(Anacardiaceae), and Citrus spp. (Rutaceae). Thefrequency of records for these species accounted
for slightly more than 30% of the total number of reports. Thepeach is recognized as the exotic
host that is most frequently reported throughout the distributional range of the SAFF. However, at
a regional level, there is a higher proportion of records for other species, such as coffee, mango,
and citrus because each of these have economic signicance in specic regions or countries
(Figure 6.2). For instance, infestation of Coffea arabica coffee cherries was found mainly in
FIGURE6.1 Host plant families for the Anastrepha fraterculus complex recorded in the Americas. Bars
indicate the number of plant species; black bars, native species; white bars, exotic species.
110 Area-Wide Management of Fruit Fly Pests
FIGURE6.2 Trophic network interactions of the Anastrepha fraterculus complex with exotic host plants
recorded in various countries across the Americas. Plant species with less than ve records are omitted.
Thethickness of the linking gray lines represents the frequency of records. Same color nodes (left) denote that
the hosts belong to the same family; the red nodes (right) depict countries of distribution.
111AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
the northern region of the Andes in Venezuela and Colombia but also in some Brazilian loca-
tions. InCitrus spp., almost all of the records were from South America, especially Brazil and
Argentina, whereas fruits of Mangifera indica have been most commonly reported as hosts in
Brazil, Argentina, and Ecuador. These results revealed the occurrence of certain preferences in the
use of hosts at a regional level, suggesting the relatively recent adaptation of some y populations
in order to exploit those exotic hosts.
On the other hand, the list of native hosts comprised a total of 124 plant species throughout the
American tropics, accounting for 70% of all of the known hosts listed. Thehighest frequencies of
records were found for Psidium guajava (298), Eugenia uniora (48), Acca sellowiana (31), Spondias
purpurea (23), Psidium cattleianum (22), and Annona cherimola (21), among others. Itshould be
noted that 8 of the 10 native species with the highest number of records belonged to the family
Myrtaceae, which represented nearly 29% of the total. Itis also noteworthy that guava fruits are the
only common host shared by different populations throughout the tropical and subtropical regions of
the Americas. Incontrast, other common host species (e.g., Surinam cherry, feijoa, seriguela, araçá,
and chirimoya) are only found infested in certain biogeographical provinces of the neotropical region
(Figure6.3).
6.3.3 geographICal dIstrIbutIon
Examination of the geographical distribution of host plants used by the SAFF produced records
for 12Latin American countries. Thelargest numbers of host plants were documented for South
America, including Brazil (121), Argentina (40), Ecuador (40), Colombia (38), and Venezuela (24).
InMexico, one of the most well-studied countries in the north of the Americas, the infestation of
19host plants has been recorded.
The data on host plants recorded for each country allowed us to explore shared resources
through a cluster analysis. In this context, Brazil exhibited the highest dissimilarities of hosts
shared with all other countries. Thisis supported by previous studies that highlight the presence
of three morphotypes in this territory (Selivon 2004, 2005a, 2005b), which would explain the
existence of 120 host species, including many native species. Incontrast, Colombia, Ecuador, and
Argentina added another subgroup with closer afnities in terms of resource use. These countries
shared exotic plants such as Prunus domestica, Punica granatum, and Ficus carica, but also native
plants such as Acca sellowiana, Malphigia glabra, and Annona cherimola, among others. Mexico
and Venezuela formed a third cluster, sharing hosts such as Terminalia catappa, Syzygium jambos,
and Psidium guineense (Figure6.4).
With respect to taxonomic richness, 30 plant families were found in Brazil, representing nearly
twice the number of families found in other countries. Thiscontrasts with the situation in Argentina,
where only 16 families were recognized. TheBrazilian territory is larger, and three distinct mor-
photypes have been characterized within the SAFF species complex, whereas just one of them has
been reported in Argentina (Hernández-Ortiz etal. 2012). Thishas led to the hypothesis that the
widespread host range found in Brazilian populations could be explained by the presence of differ-
ent taxa, which requires further investigation.
Therichness in the Andean region comprised 10–17families and 24–40plant species. Those
countries share the common infestation of coffee berries, pineapple guava, custard apple, and berry
of the Andes, among others. These are hosts that are rare or absent elsewhere in the Americas.
In the entire northern region, from Mesoamerica to Central America, including Mexico, Costa
Rica, Guatemala, and Panama, 33 host plant species are recognized. In the northern and central
Andean countries, such as Venezuela, Colombia, Ecuador, Peru, and Bolivia, a total of 68 host
plants are known. Incontrast, in the Amazonian, Atlantic Forest and Chacoan subregions of Brazil
and Argentina, a total of 135 plant species are recorded.
112 Area-Wide Management of Fruit Fly Pests
FIGURE6.3 Network of trophic interactions of the Anastrepha fraterculus complex with native host plants
recorded in various countries across the Americas. Plant species with less than ve records are omitted.
Thethickness of the linking gray lines represents the frequency of records. Same color nodes (left) denote that
the hosts belong to the same family; the red nodes (right) depict countries of distribution.
113AReview of the Natural Host Plants of the Anastrepha fraterculus Complex in the Americas
6.4 REMARKS AND CONCLUSIONS
Thecurrent knowledge of host plants occupied by populations of the nominal species A. fratercu-
lus throughout the Americas shows the persistence of gaps in the information. Thiswas evident in
nearly all of the countries of Central America and some others from South America (i.e., Bolivia,
Peru, Surinam, Paraguay, and Uruguay).
Plant richness showed a great diversication of hosts under natural conditions constituted by
70.1% of native species. Prominent among these are the species of the Myrtaceae family, which
accounted for 25% of the total number of host species. These plants could therefore have been
the most important inuence in the original diversication of the A. fraterculus species complex.
However, other families, such as the Sapotaceae, Annonaceae, Leguminosae, Anacardiaceae,
Passioraceae, and Solanaceae, might have been used secondarily. Theexotic species belonged to
17botanical families, although most were represented only by a few species. Thenotable exceptions
to this were the two richest families, Rosaceae and Rutaceae, which together accounted for two-
thirds of all the exotic species recorded, particularly of plants within the respective genera Prunus
and Citrus.
Thehigh richness of native host plants for the A. fraterculus complex denotes an explicit sign of
host diversication. However, it should be noted that exotic plants have played a signicant role in
the divergence of the species complex because some have become almost exclusive hosts of popu-
lations in certain biogeographical zones. Forexample, in the central region of the Andes, Coffea
arabica is widely used as a preferred host by the SAFF but is virtually an unknown host in other
neotropical locations. Thissuggests that such plants could have played a critical role in the current
dispersion of the SAFF following their introduction to the Americas.
Theareas occupied by certain populations of this nominal species reect some preferences in the
use of resources. Thiswould reinforce the hypothesis regarding the existence of distinct taxonomic
entities because, although these resources exist throughout the range, they are notcommon hosts
FIGURE 6.4 Dendrogram of dissimilarities (Ward distance: 0–2.33) by countries across the Americas,
based on qualitative records (presence–absence)of the host plant species.
114 Area-Wide Management of Fruit Fly Pests
for all y populations. Thefact that there are at least eight morphotypes, presumably corresponding
to different species within the SAFF complex, leads us to conclude that knowledge on host use at a
regional level is critical. Thedata analyzed here indicated strong preferences at the regional level for
both native and introduced plants, and the latter have served as a scattering factor, thus converting
some of these populations into distinct species or hosting races of economic importance.
Inthis context, the potential application of the SIT in certain geographical areas requires
knowledge in terms of the host plants consumed by the target species. Aholistic view of the
trophic interactions among populations of this complex allows the use of much of the published
host information. Thiscould help to outline a condence range of preferred hosts exploited by
each biological species and ultimately determine their pest status in different regions of the
Americas.
ACKNOWLEDGMENTS
Thisresearch received nancial support from the International Atomic Energy Agency (Vienna,
Austria) through the Research Contract No:22457/R0–MEX. We thank José F. Dzul-Cauich for
technical assistance with capturing of part of the databases. We appreciate the feedback and sugges-
tions from two anonymous reviewers that improved the nal version of the manuscript.
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