Biota Neotropica 23(4): e20231568, 2023

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ISSN 1676-0611 (online edition)

Inventory

Occurrence and characterization of insect galls in two reserves of

the Peruvian Amazon

Julio Miguel Grandez-Rios1* , Walter Santos de Araújo2, Eurídice N. Honorio Coronado3,

Timothy R. Baker4, Rodolfo Vásquez Martínez5, Abel Monteagudo Mendoza5,

Roosevelt García-Villacorta6, Gerardo Flores Llampazo7, José Reyna Huaymacari7,

Maria Antonieta Pereira de Azevedo1 & Valeria Cid Maia1

1
Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, São Cristóvão,
20940-040, Rio de Janeiro, RJ, Brasil.
2
Universidade Estadual de Montes Claros, Departamento de Biologia Geral, Montes Claros, MG, Brasil.
3
University of St Andrews, School of Geography and Sustainable Development, St Andrews, United Kingdom.
4
University of Leeds, School of Geography, LS2 9JT, Leeds, United Kingdom.
5
Jardín Botánico de Missouri, Oxapampa, Peru.
6
Centro de Innovación Científica Amazónica, Programa Restauración de Ecosistemas, Puerto Maldonado, Peru.
7
Instituto de Investigaciones de la Amazonia Peruana, Iquitos, Peru.
*Corresponding author: jgrandezrios.86@gmail.com

GRANDEZ-RIOS, J.M., ARAÚJO, W.S., CORONADO, E.N.H., BAKER, T.R., VÁSQUEZ MARTÍNEZ, R.,
MONTEAGUDO MENDOZA, A., GARCÍA-VILLACORTA, R., FLORES LLAMPAZO, G., REYNA
HUAYMACARI, J., AZEVEDO, M.A.P., MAIA, V.C. Occurrence and characterization of insect galls in two
reserves of the Peruvian Amazon. Biota Neotropica 23(4): e20231568. https://doi.org/10.1590/1676-0611-
BN-2023-1568

Abstract: An insect gall inventory was carried out in two reserves of the Peruvian Amazon, Allpahuayo-Mishana
National Reserve and Quistococha Regional Reserve, both situated in Iquitos, northeastern Peru. Four vegetation
types were surveyed between December, 2021 and December, 2022: terra firme forest, white-sand wet forest, and
white-sand dry forest in Allpahuayo-Mishana National Reserve, and palm swamp forest in Quistococha Regional
Reserve. Overall, we found 262 gall morphotypes, distributed across 75 host species representing 66 plant genera
and 30 families. Fabaceae was the plant family with the greatest number of gall morphotypes (n = 48), followed
by Calophyllaceae (n = 21) and Euphorbiaceae (n = 20). The plant genera that supported the highest diversity of
galls were Caraipa (n = 17), Eschweilera (n = 16), Tapirira (n = 16), Micrandra (n = 14), and Iryanthera (n = 10).
The plant species Tapirira guianensis (n = 16), Caraipa utilis (n = 14), Micrandra elata (n = 14), Eschweilera
coriacea (n = 11), and Sloanea parvifructa (n = 10) exhibited the highest richness of galls. Among the host plants,
C. utilis stands alone as the only species noted as both endemic to the Amazonian region and bearing a Vulnerable
(VU) conservation status. The leaves were the most attacked organs (90% of all galls). Most morphotypes are
glabrous (89%), green (67%), globoid (53%), and one-chambered (91%). We found galling insects belonging to
the orders Diptera, Thysanoptera, Lepidoptera, and Hemiptera. The galling insects of Cecidomyiidae (Diptera)
were the most common, inducing 22% of the gall morphotypes. In addition to the gallers, we also observed the
presence of successors, cecidophages, and parasitoids. Among the sampled vegetation types, the terra firme forest
presented the highest richness of gall morphotypes and host plant species. This is the first systematic inventory of
insect galls in this part of the Peruvian Amazon.
Keywords: Cecidomyiidae; Fabaceae; galls; host plant.

Ocorrência e caracterização de galhas de insetos em duas reservas da Amazônia peruana

Resumo: Um inventário de galhas de insetos foi realizado em duas reservas da Amazônia peruana, Reserva Nacional
Allpahuayo-Mishana e Reserva Regional Quistococha, ambas situadas em Iquitos, nordeste do Peru. Quatro tipos
de vegetação foram pesquisados entre dezembro de 2021 e dezembro de 2022: floresta de terra firme, floresta úmida
de areia branca e floresta seca de areia branca na Reserva Nacional Allpahuayo-Mishana, e floresta de pântano de
palmeiras na Reserva Regional Quistococha. No total, encontramos 262 morfotipos de galhas, distribuídos em 75
espécies hospedeiras representando 66 gêneros de plantas e 30 famílias. Fabaceae foi a família de plantas com o

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Grandez-Rios J.M. et al.

maior número de morfotipos de galhas (n = 48), seguida por Calophyllaceae (n = 21) e Euphorbiaceae (n = 20). Os
gêneros de plantas que apresentaram a maior diversidade de galhas foram Caraipa (n = 17), Eschweilera (n = 16),
Tapirira (n = 16), Micrandra (n = 14) e Iryanthera (n = 10). As espécies de plantas Tapirira guianensis (n = 16),
Caraipa utilis (n = 14), Micrandra elata (n = 14), Eschweilera coriacea (n = 11) e Sloanea parvifructa (n = 10)
apresentaram a maior riqueza de galhas. Dentre as plantas hospedeiras, C. utilis destaca-se como a única espécie
listada como endêmica da região amazônica e com um status de conservação Vulnerável (VU). As folhas foram os
órgãos mais atacados (90% de todas as galhas). A maioria dos morfotipos é glabra (89%), verde (67%), globoide
(53%) e possui apenas uma câmara interna (91%). Encontramos insetos galhadores pertencentes às ordens Diptera,
Thysanoptera, Lepidoptera e Hemiptera. Os insetos galhadores da família Cecidomyiidae (Diptera) foram os
mais comuns, induzindo 22% dos morfotipos de galhas. Além dos galhadores, também observamos a presença de
sucessores, cecidófagos e parasitoides. Entre os tipos de vegetação amostrados, a floresta de terra firme apresentou
a maior riqueza de morfotipos de galhas e espécies de plantas hospedeiras. Este é o primeiro inventário sistemático
de galhas de insetos nesta região da Amazônia peruana.
Palavras-chave: Cecidomyiidae; Fabaceae; galha; planta hospedeira.

Introduction firme forest lies in the unique characteristics of this habitat (e.g., plant
diversity and soil fertility) (Julião et al. 2014b). Furthermore, terra
Peru is a mega-diverse country in terms of species richness, and firme forest exhibits a remarkable diversity array of plants, providing
comprises a large part of the Amazon forest, which constitutes over 70% a wide range availability of ecological niches and an abundant source
of the national territory (Gagliardi-Urrutia et al. 2022). Estimates based of resources when compared to the more limited environments, such
on floristic inventories in the Peruvian Amazon suggest that there are as white-sand and palm swamp forests (Draper et al. 2018). In this
approximately 5,000 plant species (Cardoso et al. 2017), almost 756 context, the terra firme forest is expected to have a higher diversity of
(13%) of which are endemic (Leon et al. 2006). This high number of gall-inducing insect than other Amazonian vegetation.
plant species represents a great diversity of potential niches for gall- Over the last 30 years, several insect gall inventories have been
inducing insects (Mendonça 2007). Galling insects are very specialized carried out in different phytogeographic domains of the Neotropics,
herbivores and tend to be species-specific with respect to host plants including in the Amazon, although it is one of the least studied (Araújo
(Stone & Schönrogge 2003, Carneiro et al. 2009). These insects are 2018). In fact, there is a significant bias for studies in Southeast Brazil,
also very diverse, with some estimates indicating that there could be where the most important Brazilian research centers regarding galls
as many as 132,000 species worldwide (Espírito-Santo & Fernandes and galling insects are located, as shown by Araújo (2018). Araújo et
2007), and 3,090 in Peru (Grandez-Rios et al. 2015). al. (2019) compiled data of four inventories performed in the Brazilian
Taxonomic knowledge of gall-inducing insects in the Amazon forest Amazon. Despite this small number of published inventories, the galling
is scarce, largely because of taxonomic uncertainties and difficulty species richness was greater than in other Brazilian biomes, such as
of identification, or else because the species have yet to be described Cerrado. These results show that Amazonian forests host a high galling
(review in Grandez-Rios et al. 2015). Most inventories of galling insects insect diversity, likely due to the high specialization associated with
therefore identify taxa only to the order or family level (Yukawa et al. the remarkable richness of host plants. However, in other parts of the
2001, Almada & Fernandes 2011), or rarely to genus (Maia 2011). Galling Amazon, such as Peru, insect gall inventories are lacking, and little is
insects recorded in the Amazon forest belong to Diptera, Hemiptera, known about host plant associations. The objective of the present study
Coleoptera, Lepidoptera and Hymenoptera, with the Dipteran family was to inventory the galling insects, associated fauna, and their host
Cecidomyiidae particularly well represented (Julião et al. 2014b). plants in the Peruvian Amazon.
Cecidomyiidae comprise more than 6,500 described species worldwide
(Gagné & Jaschhof 2021), and achieves its greatest diversity in Amazonia Materials and Methods
(Grandez-Rios et al. 2015). Insect gall inventories in Amazonia have
found that Fabaceae is the most important host plant family (Almada 1. Study area
& Fernandes 2011, Maia 2011, Silva et al. 2011, Araújo et al. 2012,
Julião et al. 2017). Other host plant families include Burseraceae, The study was realized southwest of the city of Iquitos, in the
Chrysobalanaceae, Sapotaceae and Lecythidaceae (Julião et al. 2014b). province of Loreto in the Peruvian Amazon. The area exhibits a
Each of these families is notably species-rich in Amazonian forests. warm and moist climate with an average annual temperature 26 °C
Studies carried out in the Amazonian region have also show that and average annual precipitation of 3,087 mm (Marengo 1998). The
the diversity of gall-inducing insects is highest in terra firme forests site is characterized by a long wet season with monthly precipitations
in comparison with other vegetation types (Julião et al. 2014a). The > 300 mm in November, March and April, and a weak dry season in June
reason behind this greater diversity of gall-inducing insects in terra and September. (Marengo 1998). Humidity is 80–90% all year round.

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Insect galls of the Peruvian Amazon

The research was conducted in three vegetation types in the protocol (Phillips et al. 2009). Altogether 32 plots were established,
Allpahuayo-Mishana National Reserve (RNAM): terra firme forest being 24 plots for RNAM (8 plots in terra firme forest; 8 plots in white
(S 03° 57.056′, W 073° 26.002′; S 03° 57.051′, W 073° 26.320′; sand wet forest and 8 plots in white sand dry forest) and 8 plots for
TF), white-sand wet forest (S 03° 57.062′ W 073° 24.586′; WSw) RRQ (all in palm swamp forest). The woody plant species sampled
and white-sand dry forest (S 03° 57.253′, W 073° 25.617′; WSd). In in the plots were already identified in species as part of a previous
addition, one vegetation type was assessed in the Quistococha Regional floristic project, when each individual received a permanent metal
Reserve (RRQ): palm swamp forest (S 03° 50.037′, W 073° 19.254′; tag, whose related data were downloaded via ForestPlots.net online
S 03° 50.082′, W 073° 19.019′; PS) (Figure 1). The Allpahuayo- database (Lopez-Gonzalez et al. 2011, ForestPlots.net et al. 2021).
Mishana National Reserve (58,069.90 ha.), exhibits a high degree of Data on the conservation status and endemism of each host plant
both endemism and biodiversity, largely owing to a complex mosaic of species were retrieved from the IUCN Red List of Threatened Species
habitats—ranging from white sand forest patch growing on nutrient-poor (IUCN 2023) and Leon et al. (2006).
soils, to seasonally-inundated floodplain forests, and terra firme forests All the plots were established in the understory vegetation and
growing on nutrient-rich clay soils (Salo & Pyhälä 2007). Meanwhile, then visually projected onto the canopy, delimiting the sampling area
Quistococha Regional Reserve (369 ha), is characterized by palm swamp to collect the tree crown above each individual. Ten terminal units
forests, with poorly drained soils that are often flooded (Encarnación of the plant were randomly collected by climbing, and with a 10 m
1985). These areas are dominated by the palms Mauritia flexuosa and telescoping aluminum pole pruner. In the field, all gall morphotypes
Mauritiella armata (both Arecaceae), and by the broadleaf tree Tabebuia were characterized with respect to external morphology, and by shape,
insignis (Bignoniaceae) (Roucoux et al. 2013). color, presence of trichomes, number of chambers, and the host plant
organ attacked, as proposed by Isaias et al. (2014).
2. Sampling of galling insects and host plants
Galled branches were photographed, collected and transported to
Sampling in the different vegetation types was done bi-monthly the Natural Products Chemistry Laboratory of the Peruvian Amazon
between December 2021 and December 2022, with 8 plots of 5 × 20 m Research Institute (IIAP) in closed and labeled transparent plastic
each, separated from one another by 20 m, totaling 800 m2 sampled bags. Some specimens of each gall morphotype were dissected using
for each site, in accord with the methodology proposed by Julião et al. a stylet under a Leica Wild M3C stereoscopic microscope, to observe
(2014a). In each plot, only woody plants with a diameter at breast the number of internal chambers and obtain the immature insects
height (dbh, 1.3 m) equal to or greater than 10 cm were measured (larvae and pupae). These immatures were initially preserved in 70%
and collected, using the standard methods described in the RAINFOR alcohol in labeled microtubes. The remainder of the sample was used

Figure 1. Map of the distribution of sampling areas in four different types of vegetation in the National Reserve Allpahuayo-Mishana and Regional Reserve
Quistococha, Iquitos, Peru.

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for rearing galling insects and associated fauna (successor, inquilines, Table 1. Number of host plant species and insect gall morphotypes per host plant
cecidophages, and parasitoids). To distinguish these guilds in this study, family in the Allpahuayo-Mishana National Reserve and Quistococha Regional
Reserve, Iquitos, Peru.
we used the interaction parameters proposed by Luz & Mendonça
Júnior (2019). Successors are organisms that utilize the gall after the Host family Number of Number of Number
inducer departs (Mani 1964), and they use gall tissues for both food plant genera plant species of gall
and shelter. Inquilines and cecidophages utilize galls alongside the gall morphotypes
inducer. Both feed on plant tissues and consume gall tissues, but only Anacardiaceae 1 1 16
the inquilines stimulate the growth of new tissues (Luz & Mendonça Annonaceae 3 4 7
Júnior 2019). Parasitoids are organisms that closely associate with the Apocynaceae 2 2 5
host, exploiting the host’s resources, eventually leading to the host’s
Araliaceae 1 1 3
demise upon the completion of the parasitoid development (Maia 2022).
Furthermore, each gall morphotype was placed in a closed transparent Bignoniaceae 1 1 5
plastic pot, labeled and with moistened paper at the bottom. All pots Burseraceae 1 1 1
were checked daily for adult emergence. Calophyllaceae 2 3 21
All insects were preserved in 70% alcohol. The gall midges (Diptera: Chrysobalanaceae 1 1 3
Cecidomyiidae) were later mounted on slides for microscope, following
Elaeocarpaceae 1 1 10
the methodology outlined in Gagné (1994). Cecidomyiidae genera were
Euphorbiaceae 4 4 20
determined based on the keys of Gagné available for the Neotropical
region (1994). Micro-Hymenoptera were identified to the family and Fabaceae 11 12 48
genus levels using Gibson’s key (Gibson et al. 1997). Hymenoptera Humiriaceae 1 1 2
obtained only in the larval or pupal stage remained identified only to Lauraceae 5 5 17
order. All the specimens were deposited in the Diptera collection of Lecythidaceae 3 4 18
Museu Nacional (MNNRJ)/ Universidade Federal do Rio de Janeiro,
Lepidobotryaceae 1 1 1
and at the Peruvian Amazon Research Institute (IIAP). The other insects
were sent to specialists for identification. Malpighiaceae 1 1 2
Malvaceae 3 3 12
Results Melastomataceae 1 1 2
Meliaceae 1 1 3
We found 262 gall morphotypes, distributed across 30 botanical
Metteniusaceae 1 1 2
families and comprising 66 genera and 75 species (Table 1,
Moraceae 3 3 3
Figures 2–12). The average number of gall morphotypes per host
plant species was 3.5. The plant families that showed the greatest Myristicaceae 2 4 12
richness of insect galls were Fabaceae with 48 morphotypes, Myrtaceae 3 3 6
Calophyllaceae with 21, Euphorbiaceae with 20, Lecythidaceae Nyctaginaceae 1 2 5
with 18, Lauraceae with 17, Anacardiaceae with 16, Sapotaceae Olacaceae 1 1 2
with 14, Malvaceae and Myristicaceae with 12, Phyllanthaceae
Phyllanthaceae 3 3 11
with 11, and Elaeocarpaceae with 10 morphotypes. The remaining
families had less than 10 insect gall morphotypes each. The plant Salicaceae 1 1 3
genera Caraipa Aubl. (Calophyllaceae), Eschweilera Mart. ex DC. Sapindaceae 2 3 5
(Lecythidaceae), Tapirira Aubl. (Anacardiaceae), Micrandra Benth. Sapotaceae 3 4 14
(Euphorbiaceae) and Iryanthera (A. DC.) Warb. (Myristicaceae) Urticaceae 2 2 3
hosted the highest numbers of insect gall morphotypes (17, 16, 16, 14
Total 66 75 262
and 10, respectively). The most important host plant species were T.
guianensis Aubl., C. utilis Vásquez, M. elata (Didr.) Müll. Arg., E.
coriacea (DC.) S.A. Mori, and Sloanea parvifructa Steyerm., hosting
16, 14, 14, 11 and 10 insect gall morphotypes, respectively (Table Ten distinct gall shapes were found: globoid, lenticular, fusiform,
2). On the other hand, among all host plant species, only C. utilis is marginal roll, conical, clavate, amorphous, cup, pocket, and rosette. The
mentioned as endemic to the Amazonian region with a Vulnerable globoid, lenticular and fusiform shapes were the most frequent ones,
(VU) conservation status, while T. insignis (Miq.) Sandwith was with 130, 58 and 39 morphoytpes, respectively, followed by marginal
categorized as Near Threatened (NT). The rest of the plant species roll (n = 15), conical (n = 14), clavate (n = 2), and others (n = 1). The gall
were categorized as Least Concern (LC), with some not being morphotypes included green, brown, black, and yellow coloration with
classified within the IUCN list (Table 2). green color predominant (67%), followed by brown (32%). The majority
Galls were found in leaves (n = 235), stems (n = 21), buds (n = 3) and of galls were glabrous, with 232 morphotypes (89%); the remainder
fruits (n = 2), with 90% of all galls occurring on leaves. All morphotypes were hairy, with 30 morphotypes (11%). About 91% (238 morphotypes)
occurred in just one plant organ, except one morphotype that was found of galls were one-chambered, often occupied by a single larva; just 9%
in both leaves and petioles of E. coriacea. (24 morphotypes) were multi-chambered.

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Figure 2. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 1–16; Tapirira
guianensis (1. globoid-leaf, 2. conical-leaf, 3. fusiform-leaf, 4. lenticular-leaf, 5. globoid-leaf, 6. globoid-leaf (rachis), 7. fusiform-leaf, 8. globoid-leaf (petiole),
9. globoid-leaf, 10. lenticular-leaf, 11. marginal roll-leaf, 12. conical-leaf, 13. globoid-stem, 14. globoid-leaf (rachis), 15. conical-leaf (rachis), 16. fusiform-leaf),
17–18; Diclinanona tessmannii (17. globoid-leaf, 18. globoid-stem), 19–21; Guatteria megalophylla (19. conical-leaf, 20. globoid-leaf, 21. globoid-leaf), 22; Porcelia
nitidifolia (22. globoid-leaf), 23; Unonopsis spectabilis (23. globoid-leaf), 24; Aspidosperma excelsum (24. lenticular-leaf).

Figure 3. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 25–26;
Aspidosperma excelsum (25. lenticular-leaf, 26. globoid-leaf), 27–28; Parahancornia peruviana (27. lenticular-leaf, 28. fusiform-leaf), 29–31; Dendropanax
umbellatus (29. globoid-leaf, 30. lenticular-leaf, 31. fusiform-leaf), 32–36; Tabebuia insignis (32. globoid-leaf, 33. fusiform-leaf, 34. globoid-leaf, 35.
lenticular-leaf, 36. globoid-leaf), 37; Protium crassipetalum (37. marginal roll-leaf), 38–40; Caraipa tereticaulis (38. fusiform-leaf, 39. Globoid-leaf, 40.
marginal roll-leaf), 41–48; Caraipa utilis (41. lenticular-leaf, 42. globoid-stem, 43. globoid-leaf, 44. marginal roll-leaf, 45. globoid-leaf, 46. globoid-leaf,
47. globoid-leaf, 48. globoid-leaf (petiole)).

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Figure 4. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 49–54; Caraipa utilis
(49. globoid-leaf (petiole), 50. globoid-leaf, 51. globoid-stem, 52. globoid-stem, 53. globoid-leaf, 54. globoid-stem), 55–58; Haploclathra cordata (55. lenticular-
leaf, 56. globoid-leaf, 57. globoid-leaf, 58. conical-leaf), 59–61; Couepia williamsii (59. marginal roll-leaf, 60. globoid-leaf, 61. globoid-bud), 62–71; Sloanea
parvifructa (62. lenticular-leaf, 63. marginal roll-leaf, 64. globoid-leaf, 65. globoid-stem, 66. globoid-leaf, 67. fusiform-leaf, 68. fusiform-leaf, 69. globoid-leaf, 70.
globoid-leaf, 71. lenticular-leaf), 72; Alchorneopsis floribunda (72. lenticular-leaf).

Figure 5. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 73; Alchorneopsis
floribunda (73. globoid-leaf), 74; Hevea guianensis (74. globoid-leaf), 75–88; Micrandra elata (75. globoid-leaf, 76. lenticular-leaf, 77. globoid-leaf, 78. amorphous-
leaf, 79. globoid-stem, 80. marginal roll-leaf, 81. fusiform-stem, 82. globoid-leaf, 83. globoid-leaf, 84. fusiform-leaf, 85. globoid-leaf, 86. globoid-stem, 87. globoid-
leaf, 88. globoid-stem), 89–91; Nealchornea yapurensis (89. fusiform-leaf, 90. lenticular-leaf, 91. lenticular-leaf), 92–93; Abarema auriculata (92. globoid-leaf, 93.
lenticular-leaf), 94–96; Cynometra bauhiniifolia (94. fusiform-leaf, 95. marginal roll-leafl, 96. globoid-leaf).

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Figure 6. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 97; Cynometra
bauhiniifolia (97. fusiform-stem), 98–101; Dialium guianense (98. fusiform-leaf, 99. globoid-leaf, 100. lenticular-leaf, 101. conical-leaf), 102–103; Dicymbe
uaiparuensis (102. conical-leaf, 103. lenticular-leaf), 104–111; Dimorphandra macrostachya (104. globoid-leaf, 105. lenticular-leaf, 106. globoid-leaf, 107. pocket-
leaf, 108. globoid-leaf, 109. globoid-leaf, 110. globoid-leaf, 111. globoid-leaf (rachis)), 112–114; Hymenaea courbaril (112. globoid-stem, 113. conical-leaf, 114.
lenticular-leaf), 115; Inga nobilis (115. globoid-leaf), 116–119; Macrolobium limbatum (116. globoid-leaf, 117. fusiform-leaf, 118. lenticular-fruit, 119. lenticular-
leaf), 120; Macrolobium microcalyx (120. globoid-leaf).

Figure 7. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 121–125; Macrolobium
microcalyx (121. lenticular-leaf, 122. globoid-leaf, 123. conical-leaf, 124. globoid-leaf, 125. fusiform-leaf), 126–130; Parkia igneiflora (126. fusiform-leaf, 127.
cup-leaf, 128. lenticular-leaf, 129. globoid-leaf, 130. globoid-leaf), 131–136; Tachigali loretensis (131. globoid-leaf, 132. lenticular-leaf, 133. marginal roll-leaf, 134.
globoid-leaf, 135. globoid-leaf, 136. rosette-leaf), 137–139; Vatairea erythrocarpa (137. globoid-leaf, 138. globoid-leaf, 139. lenticular-leaf), 140–141; Vantanea
parviflora (140. globoid-leaf, 141. fusiform-leaf), 142–144; Aniba taubertiana (142. lenticular-leaf, 143. globoid-leaf, 144. fusiform-leaf).

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Figure 8. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 145; Aniba taubertiana
(145. globoid-bud), 146–150; Nectandra longifolia (146. lenticular-leaf, 147, globoid-leaf, 148. globoid-leaf, 149. fusiform-leaf, 150. marginal roll-leaf), 151–154;
Ocotea argyrophylla (151. globoid-leaf, 152. globoid-leaf, 153. conical-leaf, 154. fusiform-leaf), 155–156; Persea boliviensis (155. lenticular-leaf, 156. globoid-bud),
157–158; Pleurothyrium cuneifolium (157. lenticular-stem, 158. fusiform-leaf), 159; Allantoma pluriflora (159. lenticular-leaf), 160–168; Eschweilera coriacea
(160. globoid-leaf, 161. globoid-petiole/leaf, 162. globoid-stem, 163. globoid-leaf, 164. globoid-leaf, 165. globoid-leaf, 166. globoid-leaf, 167. lenticular-leaf, 168.
marginal roll-leaf).

Figure 9. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 169–170; Eschweilera
coriacea (169. globoid-leaf, 170. globoid-leaf), 171–175; Eschweilera rufifolia (171. globoid-leaf, 172, globoid-leaf, 173. lenticular-leaf, 174. lenticular-leaf, 175.
globoid-leaf), 176; Gustavia longifolia (176. fusiform-leaf), 177; Ruptiliocarpon caracolito (177. globoid-leaf), 178–179; Bunchosia argentea (178. globoid-leaf,
179. globoid-leaf), 180–184; Luheopsis hoehnei (180. globoid-leaf, 181. globoid-leaf, 182. 183. globoid-leaf, 184. fusiform-leaf), 185–188; Pachira brevipes (185.
lenticular-leaf, 186. globoid-stem, 187. globoid-leaf, 188. fusiform-stem), 189–191; Theobroma subincanum (189. globoid-leaf, 190. globoid-leaf, 191. fusiform-
leaf), 192; Mouriri vernicosa (192. lenticular-leaf).

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Biota Neotrop., 23(4): e20231568, 2023 9

Insect galls of the Peruvian Amazon

Figure 10. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 193; Mouriri
vernicosa (193. Marginal roll-leaf), 194–196; Carapa procera (194. lenticular-leaf, 195, fusiform-leaf, 196. globoid-leaf), 197–198; Emmotum floribundum (197.
fusiform-leaf, 198. lenticular-leaf), 199; Brosimum utile (199. lenticular-leaf), 200; Perebea xanthochyma (200. globoid-leaf), 201; Trymatococcus amazonicus
(201. lenticular-fruit), 202–203; Iryanthera juruensis (202. lenticular-leaf, 203. fusiform-leaf), 204–205; Iryanthera laevis (204. fusiform-leaf, 205. lenticular-leaf),
206–211; Iryanthera lancifolia (206. lenticular-leaf, 207. lenticular-leaf, 208. lenticular-leaf, 209. fusiform-leaf, 210. conical-leaf, 211. globoid-leaf), 212–213; Virola
pavonis (212. lenticular-leaf, 213. Conical-leaf), 214; Blepharocalyx eggersii (214. lenticular-leaf); 215–216; Eugenia riparia (215. globoid-leaf, 216. fusiform-leaf).

Figure 11. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 217; Eugenia
riparia (217. globoid-leaf), 218–219; Myrcia cebra (218. lenticular-leaf, 219, globoid-stem), 220–222; Neea spruceana (220. lenticular-leaf, 221. globoid-leaf,
222. lenticular-leaf), 223–224; Neea verticillata (223. globoid-leaf, 224. lenticular-leaf), 225–226; Cathedra acuminata (225. lenticular-leaf, 226. globoid-stem),
227–230; Amanoa guianensis (227. lenticular-leaf, 228. marginal roll-leaf, 229. lenticular-leaf, 230. globoid-stem), 231–232; Hieronyma oblonga (231. globoid-leaf,
232. globoid-leaf), 233–237; Richeria grandis (233. globoid-leaf, 234. globoid-leaf, 235. globoid-leaf, 236. globoid-leaf, 237. globoid-leaf), 238–240; Casearia
arborea (238. fusiform-leaf, 239. lenticular-leaf, 240. fusiform-stem).

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 10 Biota Neotrop., 23(4): e20231568, 2023

Grandez-Rios J.M. et al.

Figure 12. Galls found in four vegetation types in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. 241; Cupania
diphylla (241. conical-leaf), 242–243; Cupania latifolia (242. Marginal roll-leaf, 243, fusiform-leaf), 244–245; Talisia firma (244. fusiform-leaf, 245. lenticular-
leaf), 246–251; Chrysophyllum sanguinolentum (246. globoid-leaf, 247. fusiform-leaf, 248. globoid-leaf, 249. globoid-leaf, 250. lenticular-leaf, 251. Fusiform-leaf),
252–255; Manilkara bidentata (252. globoid-leaf, 253. globoid-stem, 254. lenticular-leaf, 255. Marginal roll-leaf), 256–258; Pouteria subrotata (256. conical-
leaf, 257. clavate-leaf, 258. globoid-leaf), 259; Pouteria torta (259. fusiform-leaf), 260; Cecropia distachya (260. globoid-leaf), 261–262; Pouroma bicolor (261.
clavate-leaf, 262. globoid-leaf).

The galling insects were distributed across four orders: Diptera (Calorileya sp.; Phylloxeroxenus sp.; Platyrileya sp. 1; Prodecatoma
(n = 58), Thysanoptera (n = 2), Lepidoptera (n = 2), and Hemiptera sp.; Tenuipetiolus sp.), Mymaridae (Polynema sp.), Torymidae
(n = 1) (Table 3). The Cecidomyiidae (Diptera) was responsible for 22% (Torymus sp. 1, 2), Platygastridae (Magellanium sp. 1, 2), Pteromalidae
of all morphotypes. In 199 (76%) of the gall morphotypes we did not find (Cerocephala sp.; Euteloidea sp.), and Scelionidae (Baryconus sp.).
the gall-inducing insect, either because the galls were empty, contained In the Allpahuayo-Mishana National Reserve, a total of 240
parasitoids, or were occupied by multiple potentially gall-inducing gall morphotypes were found across 68 plant species, representing
taxa. Eigtheen species of gall midges were identified, in two genera: 29 botanical families and 59 genera. Meanwhile, in the Quistococha
Contarinia Rondani, 1860 with five morphospecies and Bruggmanniella Regional Reserve, 22 gall morphotypes were found, distributed among
Tavares, 1909 with one morphospecies. The tribes: Asphondyliini and 8 families, 8 genera, and 8 species. These data were also analyzed based
Lasiopteridi both contained two morphospecies, while Clinodiplosini on the type of vegetation present in each reserve.
and Lopesiini had three and five morphospecies, respectively. This is The terra firme forest showed the greatest diversity of galls and
due to scarcity of material, either because one or both sexes were absent, host plants, with 111 different morphotypes found on 39 host plant
or because no immature specimens were observed. Additionally, the species, followed by white-sand wet forest, white-sand dry forest and
inducer of two gall morphotypes was identified as Plagiothrips sp. 2 palm swamp forest, which had 73, 56 and 22 morphotypes, among 16,
(Thysanoptera). 16 and 8 host plant species respectively (Table 2). In terra firme forest,
The associated fauna comprised successors, cecidophages and leaves (n = 104) were the most commonly galled plant organ; the most
parasitoids. Successors, represented by Thysanoptera (Adraneothrips common gall morphologies included the globoid shape (n = 51), green
sp., Holothrips sp. 1, and Plagiothrips sp. 1), Hymenoptera (Formicidae color (n = 70), glabrous surface (n = 94) and single internal chamber
and Apidae), Collembola and Acari, were found in 8, 8, 5 and (n = 104). In white-sand wet forest, leaves are most commonly attacked
3 morphotypes, respectively. Cecidophages, Curculionidae (Coleoptera) (n = 65), and morphologies are mostly globoid (n = 41), green (n = 52),
and Sciaridae (Diptera) were found in only one morphotype. The glabrous (n = 67) and with a single inner chamber (n = 66). In white-sand
parasitoids obtained from 55 morphotypes (21% of the total) represented dry forest: leaves (n = 45), globoid (n = 29), green (n = 35), glabrous
11 families and 23 genera of Hymenoptera: Aphelinidae (Coccobius (n = 52), and a single inner chamber (n = 48). For palm swamp forest:
sp.), Braconidae (Allorhogas sp. 1, 2), Encyrtidae (Metaphycus sp.), leaves (n = 21), globoid (n = 9), green (n = 18), glabrous (n = 19), and
Eulophidae (Aranobroter sp.; Baryscapus sp.; Chrysonotomyia sp.; a single inner chamber (n = 20) (Table 2).
Euderus sp.; Euderomphale sp.; Galeopsomyia sp.; Horismenus sp.; The terra firme forest also showed the richest gall-inducing
Neopomphale sp. 1, 2), Eupelmidae (Lambdobregma sp.), Eurytomidae insect fauna, represented by Diptera (n = 18), Thysanoptera (n = 2),

http://www.scielo.br/bn https://doi.org/10.1590/1676-0611-BN-2023-1568
 Table 2. Characterization of insect galls in four vegetation type in the Allpahuayo-Mishana National Reserve and Quistococha Regional Reserve, Iquitos, Peru. Legends: P = Parasitoid, S = Successor,
C = Cecidophages; TF = Terra firme forest, WSd = White-sand dry forest, WSw= White-sand wet forest, and PS = Palm swamp forest.

Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Anacardiaceae Tapirira Leaf Globoid Brown Yes One-chambered TF Unidentified Least Concern (LC) Fig. 1
guianensis Aubl.
Leaf Conical Green Yes One-chambered TF Unidentified Hymenoptera (P) Fig. 2
Leaf Fusiform Brown Yes One-chambered TF Unidentified Fig. 3
Biota Neotrop., 23(4): e20231568, 2023

Leaf Lenticular Green No One-chambered TF Unidentified Fig. 4

Leaf Globoid Brown Yes One-chambered TF Unidentified Fig. 5

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Leaf Globoid Brown Yes One-chambered TF Unidentified Fig. 6
Leaf Fusiform Brown No One-chambered TF Thysanoptera Hymenoptera (P) Fig. 7
(Plagiothrips sp. 2)
Leaf Globoid Brown No One-chambered TF Thysanoptera Fig. 8
(Plagiothrips sp. 2)
Leaf Globoid Brown No One-chambered TF Cecidomyiidae Hymenoptera (P) Fig. 9
Leaf Lenticular Green No One-chambered WSd Unidentified Fig. 10
Leaf Marginal Green No One-chambered WSd Unidentified Fig. 11
roll
Leaf Conical Green No One-chambered WSd Unidentified Fig. 12
Stem Globoid Brown No One-chambered WSd Cecidomyiidae Fig. 13
Leaf Globoid Brown No One-chambered WSd Cecidomyiidae Allohorgas sp. 2 (P) Fig. 14
Insect galls of the Peruvian Amazon

(Hymenoptera)
Leaf Conical Brown No One-chambered WSd Unidentified Fig. 15
Leaf Fusiform Brown No Multilocular WSd Cecidomyiidae Hymenoptera (P) Fig. 16
Annonaceae Diclinanona Leaf Globoid Brown No One-chambered WSd Unidentified Least Concern (LC) Fig. 17
tessmannii Diels
Stem Globoid Brown No One-chambered WSd Unidentified Fig. 18
Guatteria Leaf Conical Green No One-chambered WSd Unidentified Least Concern (LC) Fig. 19
megalophylla
Diels
Leaf Globoid Green No One-chambered WSd Unidentified Formicidae (S) Fig. 20
Leaf Globoid Brown No One-chambered WSd Unidentified Fig. 21
Porcelia Leaf Globoid Green No One-chambered TF Unidentified Least Concern (LC) Fig. 22
nitidifolia Ruiz
& Pav.
Unonopsis Leaf Globoid Green Yes One-chambered TF Cecidomyiidae Least Concern (LC) Fig. 23
11

spectabilis Diels

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Apocynaceae Aspidosperma Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 24
excelsum Benth.

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Leaf Lenticular Green No One-chambered TF / WSd Cecidomyiidae Hymenoptera (P) Fig. 25
Leaf Globoid Green No One-chambered WSd Unidentified Fig. 26
Parahancornia Leaf Lenticular Green No One-chambered PS Unidentified - Fig. 27
peruviana
Monach.
Leaf Fusiform Green No One-chambered PS Unidentified Fig. 28
Araliaceae Dendropanax Leaf Globoid Brown Yes One-chambered WSw Unidentified - Fig. 29
umbellatus (Ruiz
& Pav.) J.F.
Macbr.
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 30
Leaf Fusiform Green No One-chambered WSw Cecidomyiidae Hymenoptera (P) Fig. 31
Bignoniaceae Tabebuia insignis Leaf Globoid Brown No One-chambered PS Unidentified Near Threatened Fig. 32
(Miq.) Sandwith (NT)
Leaf Fusiform Green No One-chambered PS Unidentified Polynema sp. (P) Fig. 33
(Hymenoptera)
Leaf Globoid Green No One-chambered PS Unidentified Fig. 34
Leaf Lenticular Green No One-chambered PS Unidentified Adraneothrips sp. (S) Fig. 35
Grandez-Rios J.M. et al.

(Thysanoptera)
Leaf Globoid Green No One-chambered PS Unidentified Fig. 36
Burseraceae Protium Leaf Marginal Green No One-chambered TF Cecidomyiidae Hymenoptera (P) - Fig. 37
crassipetalum roll
Cuatrec.
Holothrips sp. 1 (S)
(Thysanoptera)
Formicidae (S)
Calophyllaceae Caraipa Leaf Fusiform Green No Multilocular WSw Cecidomyiidae Hymenoptera (P) Least Concern (LC) Fig. 38
tereticaulis Tul.
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 39
Leaf Marginal Green No One-chambered WSw Cecidomyiidae Hymenoptera (P) Fig. 40
roll
Caraipa utilis Leaf Lenticular Green No One-chambered WSw Unidentified Vulnerable (VU) Fig. 41
Vásquez
Stem Globoid Brown No One-chambered WSw Unidentified Fig. 42

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Leaf Globoid Green No Multilocular WSw Cecidomyiidae Magellanium sp. 2 (P) Fig. 43
(Hymenoptera)
Holothrips sp. 1 (S)
(Thysanoptera)
Leaf Marginal Green No One-chambered WSw Cecidomyiidae Holothrips sp. 1 (S) Fig. 44
Biota Neotrop., 23(4): e20231568, 2023

roll
(Thysanoptera)
Collembola (S)

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Leaf Globoid Green No One-chambered WSw Unidentified Fig. 45
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 46
Leaf Globoid Green No One-chambered WSw Cecidomyiidae Collembola (S) Fig. 47
Formicidae (S)
Eupelmidae (P)
Braconidae (P)
(Hymenoptera)
Leaf Globoid Brown No One-chambered WSw Cecidomyiidae Tenuipetiolus sp. (P) Fig. 48
(Hymenoptera)
Leaf Globoid Brown No One-chambered WSw Unidentified Phylloxeroxenus sp. Fig. 49
(P)
Insect galls of the Peruvian Amazon

(Hymenoptera)
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 50
Stem Globoid Brown No One-chambered WSw Unidentified Fig. 51
Stem Globoid Brown No One-chambered WSw Unidentified Fig. 52
Leaf Globoid Green No Multilocular WSw Unidentified Fig. 53
Stem Globoid Brown No One-chambered WSw Cecidomyiidae Curculionidae (C) Fig. 54
(Coleoptera)
Haploclathra Leaf Lenticular Green No One-chambered WSw Unidentified - Fig. 55
cordata Vásquez
Leaf Globoid Green No Multilocular WSw Unidentified Hymenoptera (P) Fig. 56
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 57
Leaf Conical Green No One-chambered WSw Unidentified Fig. 58
Chrysobalan- Couepia Leaf Marginal Green No One-chambered WSd Lopesiini - Fig. 59
aceae williamsii J.F. roll
Macbr.
13

Leaf Globoid Green No One-chambered WSd Unidentified Hymenoptera (P) Fig. 60

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Bud Globoid Green No One-chambered WSd Contarinia sp. Fig. 61
Elaeocarpaceae Sloanea parvi- Leaf Lenticular Green No One-chambered WSw Unidentified - Fig. 62

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fructa Steyerm.
Leaf Marginal Green No One-chambered WSw Cecidomyiidae Horismenus sp. (P) Fig. 63
roll
(Hymenoptera)
Leaf Globoid Green No Multilocular WSw Lopesiini Hymenoptera (P) Fig. 64
Stem Globoid Green No Multilocular WSw Asphondyliini Collembola (S) Fig. 65
Prodecatoma sp. (P)
Platyrileya sp. 1 (P)
Torymus sp. 1, 2 (P)
Cerocephala sp. (P)
Lambdobregma sp. (P)
Allorhogas sp. 1 (P)
(Hymenoptera)
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 66
Leaf Fusiform Green No One-chambered WSw Unidentified Fig. 67
Leaf Fusiform Brown No One-chambered WSw Unidentified Fig. 68
Leaf Globoid Green No One-chambered WSw Unidentified Apidae (S) Fig. 69
Grandez-Rios J.M. et al.

(Hymenoptera)
Leaf Globoid Brown No One-chambered WSw Unidentified Fig. 70
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 71
Euphorbiaceae Alchorneopsis Leaf Lenticular Green No One-chambered PS Unidentified Least Concern (LC) Fig. 72
floribunda
(Benth.) Müll.
Arg.
Leaf Globoid Green Yes Multilocular PS Unidentified Hymenoptera (P) Fig. 73
Hevea guianensis Leaf Globoid Brown No One-chambered WSw Unidentified Least Concern (LC) Fig. 74
Aubl.
Micrandra elata Leaf Globoid Green No Multilocular WSd Unidentified Eulophidae (P) Least Concern (LC) Fig. 75
(Didr.) Müll. Arg.
(Hymenoptera)

Leaf Lenticular Green No One-chambered WSd Unidentified Fig. 76

Leaf Globoid Green No Multilocular WSd Lasiopteridi Neopomphale sp. 2 (P) Fig. 77
Calorileya sp. (P)

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Coccobius sp. (P)
(Hymenoptera)
Leaf Amorphous Green No One-chambered WSd Cecidomyiidae Neopomphale sp. 2 (P) Fig. 78
(Hymenoptera)
Stem Globoid Brown No One-chambered WSd Cecidomyiidae Hymenoptera (P) Fig. 79
Biota Neotrop., 23(4): e20231568, 2023

Leaf Marginal Green No One-chambered WSd Lopesiini Hymenoptera (P) Fig. 80

roll

https://doi.org/10.1590/1676-0611-BN-2023-1568
Plagiothrips sp.1 (S)
(Thysanoptera)
Stem Fusiform Green No One-chambered WSd Cecidomyiidae Collembola (S) Fig. 81
Formicidae (S)
Magellanium sp.1 (P)
(Hymenoptera)
Leaf Globoid Green No One-chambered WSd Cecidomyiidae Hymenoptera (P) Fig. 82
Collembola (S)
Leaf Globoid Green No One-chambered WSd Unidentified Fig. 83
Leaf Fusiform Green No One-chambered WSd Unidentified Fig. 84
Leaf Globoid Brown No One-chambered WSd Unidentified Fig. 85
Insect galls of the Peruvian Amazon

Stem Globoid Brown No One-chambered WSd Cecidomyiidae Hymenoptera (P) Fig. 86

Leaf Globoid Green No One-chambered WSd Unidentified Fig. 87
Stem Globoid Brown No One-chambered WSd Unidentified Fig. 88
Nealchornea Leaf Fusiform Green No One-chambered TF Unidentified Least Concern (LC) Fig. 89
yapurensis
Huber
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 90
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 91
Fabaceae Abarema Leaf Globoid Green Yes One-chambered WSw Lopesiini Hymenoptera (P) Least Concern (LC) Fig. 92
auriculata
(Benth.) Barneby
& J.W. Grimes
Leaf Lenticular Green No One-chambered WSw Cecidomyiidae Fig. 93
Cynometra Leaf Fusiform Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 94
bauhiniifolia
Benth.
Leaf Marginal Green No One-chambered TF Unidentified Fig. 95
15

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Leaf Globoid Green No One-chambered TF Lasiopteridi Acari (S) Fig. 96
Stem Fusiform Brown No One-chambered TF Unidentified Fig. 97

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Dialium Leaf Fusiform Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 98
guianense
(Aubl.) Sandwith
Leaf Globoid Green No One-chambered TF Clinodiplosini Hymenoptera (P) Fig. 99
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 100
Leaf Conical Green Yes One-chambered TF Cecidomyiidae Fig. 101
Dicymbe Leaf Conical Green Yes One-chambered WSw Cecidomyiidae Baryscapus sp. (P) - Fig. 102
uaiparuensis
R.S. Cowan
(Hymenoptera)
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 103
Dimorphandra Leaf Globoid Green No One-chambered WSw Unidentified Least Concern (LC) Fig. 104
macrostachya
Benth.
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 105
Leaf Globoid Green Yes One-chambered WSw Cecidomyiidae Hymenoptera (P) Fig. 106
Leaf Pocket Green No One-chambered WSw Cecidomyiidae Galeopsomyia sp. (P) Fig. 107
(Hymenoptera)
Grandez-Rios J.M. et al.

Leaf Globoid Green No One-chambered WSw Clinodiplosini Hymenoptera (P) Fig. 108
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 109
Leaf Globoid Green No One-chambered WSw Unidentified Sciaridae (C) Fig. 110
(Diptera)
Leaf Globoid Brown No One-chambered WSw Unidentified Fig. 111
Hymenaea Stem Globoid Brown No Multilocular TF Asphondyliini Euteloidea sp. (P) Least Concern (LC) Fig. 112
courbaril L.
(Hymenoptera)
Leaf Conical Green No Multilocular TF Bruggmanniella sp. Fig. 113
Leaf Lenticular Brown No One-chambered TF Unidentified Fig. 114
Inga nobilis Leaf Globoid Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 115
Willd.
Macrolobium Leaf Globoid Green No One-chambered WSd Unidentified Least Concern (LC) Fig. 116
limbatum Spruce
ex Benth.
Leaf Fusiform Brown No One-chambered WSd Unidentified Fig. 117

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Fruit Lenticular Brown No One-chambered WSd Lepidoptera Fig. 118
Leaf Lenticular Green No One-chambered WSd Unidentified Fig. 119
Macrolobium Leaf Globoid Green No Multilocular WSd Unidentified Least Concern (LC) Fig. 120
microcalyx Ducke
Leaf Lenticular Green No One-chambered WSd / Unidentified Fig. 121
Biota Neotrop., 23(4): e20231568, 2023

WSw
Leaf Globoid Green No Multilocular WSw Unidentified Fig. 122

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Leaf Conical Green No One-chambered WSw Unidentified Fig. 123
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 124
Leaf Fusiform Brown No One-chambered WSw Unidentified Fig. 125
Parkia igneiflora Leaf Fusiform Green No One-chambered WSw Unidentified Least Concern (LC) Fig. 126
Ducke
Leaf Cup Yellow Yes One-chambered WSw Cecidomyiidae Braconidae (P) Fig. 127
(Hymenoptera)
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 128
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 129
Leaf Globoid Green No One-chambered WSw Unidentified Hymenoptera (P) Fig. 130
Tachigali loreten- Leaf Globoid Green No One-chambered WSd Unidentified - Fig. 131
sis van der Werff
Insect galls of the Peruvian Amazon

Leaf Lenticular Green No One-chambered WSd Unidentified Fig. 132

Leaf Marginal Green Yes One-chambered WSd Unidentified Formicidae (S) Fig. 133
roll
Holothrips sp. 1 (S)
(Thysanoptera)
Leaf Globoid Green No One-chambered WSd Unidentified Fig. 134
Leaf Globoid Green Yes Multilocular WSd Unidentified Fig. 135
Leaf Rosette Green Yes Multilocular WSd Cecidomyiidae Hymenoptera (P) Fig. 136
Vatairea Leaf Globoid Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 137
erythrocarpa
(Ducke) Ducke
Leaf Globoid Brown No One-chambered TF Unidentified Fig. 138
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 139
Humiriaceae Vantanea Leaf Globoid Green No One-chambered TF Unidentified Least Concern (LC) Fig. 140
parviflora Lam.
17

Leaf Fusiform Green No One-chambered TF Lepidoptera Fig. 141

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Lauraceae Aniba Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 142
taubertiana Mez

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Leaf Globoid Green No One-chambered TF Unidentified Fig. 143
Leaf Fusiform Green No One-chambered TF Unidentified Fig. 144
Bud Globoid Brown Yes One-chambered TF Unidentified Fig. 145
Nectandra Leaf Lenticular Green No One-chambered TF Unidentified - Fig. 146
longifolia (Ruiz
& Pav.) Nees
Leaf Globoid Green No One-chambered TF Unidentified Fig. 147
Leaf Globoid Green No One-chambered TF Cecidomyiidae Fig. 148
Leaf Fusiform Brown No One-chambered TF Unidentified Fig. 149
Leaf Marginal Green No One-chambered TF Unidentified Fig. 150
roll
Ocotea Leaf Globoid Brown Yes One-chambered WSd Unidentified Least Concern (LC) Fig. 151
argyrophylla
Ducke
Leaf Globoid Brown No One-chambered WSd Unidentified Hymenoptera (P) Fig. 152
Leaf Conical Brown No Multilocular WSd Unidentified Fig. 153
Leaf Fusiform Brown No One-chambered WSd Unidentified Fig. 154
Persea boliviensis Leaf Lenticular Green No One-chambered WSd Unidentified - Fig. 155
Grandez-Rios J.M. et al.

Mez & Rusby ex

Rusby
Bud Globoid Green No One-chambered WSd Cecidomyiidae Fig. 156
Pleurothyrium Leaf Lenticular Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 157
cuneifolium
Nees
Leaf Fusiform Brown No One-chambered TF Unidentified Fig. 158
Lecythidaceae Allantoma Leaf Lenticular Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 159
pluriflora S.A.
Mori, Y.Y. Huang
& Prance
Eschweilera Leaf Globoid Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 160
coriacea (DC.)
S.A. Mori
Leaf- Globoid Brown No One-chambered TF Cecidomyiidae Holothrips sp. 1 (S) Fig. 161
Petiole
(Thysanoptera)

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Stem Globoid Brown No One-chambered TF Cecidomyiidae Fig. 162
Leaf Globoid Green No One-chambered TF Contarinia sp. Formicidae (S) Fig. 163
Baryconus sp. (P)
(Hymenoptera)
Leaf Globoid Brown No One-chambered TF Unidentified Fig. 164
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Leaf Globoid Green No One-chambered TF Unidentified Fig. 165

Leaf Globoid Green No Multilocular TF Cecidomyiidae Fig. 166

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Leaf Lenticular Green No One-chambered TF Unidentified Fig. 167
Leaf Marginal Green No One-chambered TF Unidentified Fig. 168
roll
Leaf Globoid Brown No One-chambered TF Unidentified Fig. 169
Leaf Globoid Brown No One-chambered TF Contarinia sp. Fig. 170
Eschweilera rufi- Leaf Globoid Brown No Multilocular TF Unidentified Formicidae (S) - Fig. 171
folia S.A. Mori
Plagiothrips sp.1 (S)
(Thysanoptera)
Braconidae (P)
(Hymenoptera)
Leaf Globoid Brown No One-chambered TF Unidentified Hymenoptera (P) Fig. 172
Insect galls of the Peruvian Amazon

Leaf Lenticular Brown Yes One-chambered TF Unidentified Fig. 173

Leaf Lenticular Green No One-chambered TF Unidentified Fig. 174
Leaf Globoid Green No Multilocular TF Unidentified Fig. 175
Gustavia Leaf Fusiform Brown No One-chambered TF Unidentified - Fig. 176
longifolia Poepp.
ex O. Berg
Lepidobotry- Ruptiliocarpon Leaf Globoid Green No Multilocular TF Unidentified Eupelmidae (P) Least Concern (LC) Fig. 177
aceae caracolito Ham-
mel & N. Zamora
(Hymenoptera)
Malpighiaceae Bunchosia Leaf Globoid Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 178
argentea (Jacq.)
DC.
Leaf Globoid Green No One-chambered TF Unidentified Fig. 179
Malvaceae Lueheopsis Leaf Globoid Brown Yes One-chambered PS Unidentified - Fig. 180
hoehnei Burret
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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Leaf Globoid Green Yes One-chambered PS Lopesiini Euderus sp. (P) Fig. 181
(Hymenoptera)

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Leaf Globoid Brown No One-chambered PS Unidentified Fig. 182
Leaf Globoid Green No Multilocular PS Unidentified Chrysonotomyia Fig. 183
sp. (P)
Euderomphale sp. (P)
Metaphycus sp. (P)
(Hymenoptera)
Leaf Fusiform Green No One-chambered PS Cecidomyiidae Hymenoptera (P) Fig. 184
Pachira brevipes Lenticular Green No One-chambered WSw Unidentified Least Concern (LC) Fig. 185
(A. Robyns) W.S.
Alverson
Leaf
Stem Globoid Brown No One-chambered WSw Unidentified Fig. 186
Leaf Globoid Brown No One-chambered WSw Unidentified Fig. 187
Stem Fusiform Brown No One-chambered WSw Unidentified Fig. 188
Theobroma Leaf Globoid Green Yes One-chambered TF Unidentified Least Concern (LC) Fig. 189
subincanum
Mart.
Grandez-Rios J.M. et al.

Leaf Globoid Brown Yes One-chambered TF Unidentified Fig. 190

Leaf Fusiform Brown Yes One-chambered TF Unidentified Fig. 191
Melastomataceae Mouriri Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 192
vernicosa
Naudin
Leaf Marginal Green No One-chambered TF Unidentified Hymenoptera (P) Fig. 193
roll
Meliaceae Carapa procera Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 194
DC.
Leaf Fusiform Brown No One-chambered TF Unidentified Fig. 195
Leaf Globoid Green No One-chambered TF Unidentified Fig. 196
Metteniusaceae Emmotum Leaf Fusiform Brown No One-chambered WSw Unidentified - Fig. 197
floribundum
R.A. Howard
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 198
Moraceae Brosimum utile Leaf Lenticular Green No One-chambered PS Unidentified Least Concern (LC) Fig. 199
(Kunth) Oken

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Perebea Leaf Globoid Green Yes One-chambered TF Unidentified Least Concern (LC) Fig. 200
xanthochyma H.
Karst.
Trymatococcus Fruit Lenticular Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 201
amazonicus
Poepp. & Endl.
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Myristicaceae Iryanthera Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 202
juruensis Warb.

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Leaf Fusiform Brown No One-chambered TF Unidentified Fig. 203
Iryanthera laevis Leaf Fusiform Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 204
Markgr.
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 205
Iryanthera Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 206
lancifolia Ducke
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 207
Leaf Lenticular Black Yes One-chambered TF Unidentified Fig. 208
Leaf Fusiform Green No One-chambered TF Unidentified Fig. 209
Leaf Conical Green Yes One-chambered TF Unidentified Fig. 210
Leaf Globoid Green No One-chambered TF Unidentified Fig. 211
Virola pavonis (A. Leaf Lenticular Brown No One-chambered PS / WSd Contarinia sp. Hymenoptera (P) Least Concern (LC) Fig. 212
Insect galls of the Peruvian Amazon

DC.) A.C. Sm.

Leaf Conical Green No One-chambered PS Cecidomyiidae Fig. 213
Myrtaceae Blepharocalyx Leaf Lenticular Green No One-chambered WSd Unidentified Least Concern (LC) Fig. 214
eggersii
(Kiaersk.)
Landrum
Eugenia riparia Leaf Globoid Green No One-chambered TF Unidentified Neopomphale sp. - Fig. 215
DC. 1 (P)
(Hymenoptera)
Leaf Fusiform Green No One-chambered TF Unidentified Fig. 216
Leaf Globoid Green Yes One-chambered TF Unidentified Fig. 217
Myrcia crebra Leaf Lenticular Brown No One-chambered WSd Cecidomyiidae Least Concern (LC) Fig. 218
(McVaugh) A.R.
Lourenço & E.
Lucas
Stem Globoid Brown No One-chambered WSd Unidentified Fig. 219
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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Nyctaginaceae Neea spruceana Leaf Lenticular Green No One-chambered TF Unidentified Least Concern (LC) Fig. 220
Heimerl

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Leaf Globoid Green No One-chambered TF Cecidomyiidae Fig. 221
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 222
Neea verticillata Leaf Globoid Green No One-chambered WSd Unidentified Hymenoptera (P) - Fig. 223
Ruiz & Pav.
Leaf Lenticular Green No One-chambered WSd Unidentified Fig. 224
Olacaceae Cathedra Leaf Lenticular Green No One-chambered WSd Unidentified Hymenoptera (P) Least Concern (LC) Fig. 225
acuminata
(Benth.) Miers
Stem Globoid Brown No Multilocular WSd Unidentified Fig. 226
Phyllanthaceae Amanoa Leaf Lenticular Green No One-chambered PS Unidentified Least Concern (LC) Fig. 227
guianensis Aubl.
Leaf Marginal Green No One-chambered PS Cecidomyiidae Fig. 228
roll
Leaf Lenticular Green No One-chambered PS Unidentified Fig. 229
Stem Globoid Brown No One-chambered PS Unidentified Fig. 230
Hieronyma Leaf Globoid Green No One-chambered TF Contarinia sp. Hymenoptera (P) Least Concern (LC) Fig. 231
oblonga (Tul.)
Müll. Arg.
Grandez-Rios J.M. et al.

Leaf Globoid Green No One-chambered TF Unidentified Acari (S) Fig. 232

Richeria grandis Leaf Globoid Green No One-chambered TF Unidentified - Fig. 233
Vahl
Leaf Globoid Green No Multilocular TF Unidentified Fig. 234
Leaf Globoid Green No One-chambered TF Unidentified Acari (S) Fig. 235
Leaf Globoid Green No One-chambered TF Unidentified Fig. 236
Leaf Globoid Green No One-chambered TF Hemiptera Aranobroter sp. (P) Fig. 237
(Psyllidae)
(Hymenoptera)
Salicaceae Casearia arborea Leaf Fusiform Green No One-chambered TF Unidentified Least Concern (LC) Fig. 238
(Rich.) Urb.
Leaf Lenticular Green No One-chambered TF Unidentified Fig. 239
Stem Fusiform Brown No One-chambered TF Unidentified Fig. 240

Sapindaceae Cupania diphylla Leaf Conical Brown Yes One-chambered WSw Unidentified Least Concern (LC) Fig. 241
Vahl

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Family Species Organ Shape Color Pubescence Chambers Vegetation Gall-inducers Associated fauna Conservation status Figure
type (IUCN 2023) number
Cupania latifolia Leaf Marginal Green Yes One-chambered TF Clinodiplosini Least Concern (LC) Fig. 242
Kunth roll

Leaf Fusiform Green No One-chambered TF Unidentified Fig. 243

Talisia firma Leaf Fusiform Green No One-chambered PS Unidentified Least Concern (LC) Fig. 244
Radlk.
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Leaf Lenticular Green No One-chambered PS Unidentified Fig. 245

Sapotaceae Chrysophyllum Leaf Globoid Brown No One-chambered WSw Unidentified Least Concern (LC) Fig. 246

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sanguinolentum
(Pierre) Baehni
Leaf Fusiform Brown No One-chambered WSw Unidentified Fig. 247
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 248
Leaf Globoid Green No One-chambered WSw Unidentified Fig. 249
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 250
Leaf Fusiform Green No One-chambered WSw Unidentified Fig. 251
Manilkara Leaf Globoid Green No One-chambered WSw Unidentified Hymenoptera (P) Least Concern (LC) Fig. 252
bidentata (A.
DC.) A. Chev.
Stem Globoid Brown No One-chambered WSw Cecidomyiidae Fig. 253
Leaf Lenticular Green No One-chambered WSw Unidentified Fig. 254
Insect galls of the Peruvian Amazon

Leaf Marginal Green No One-chambered WSw Unidentified Fig. 255

roll
Pouteria Leaf Conical Green No One-chambered TF Unidentified Least Concern (LC) Fig. 256
subrotata
Cronquist
Leaf Clavate Green No One-chambered TF Unidentified Fig. 257
Leaf Globoid Green No One-chambered TF Unidentified Fig. 258
Pouteria torta Leaf Fusiform Green No One-chambered TF Unidentified Least Concern (LC) Fig. 259
(Mart.) Radlk.
Urticaceae Cecropia Leaf Globoid Green No One-chambered TF Cecidomyiidae Least Concern (LC) Fig. 260
distachya Huber
Pourouma Leaf Clavate Brown No One-chambered TF Unidentified Least Concern (LC) Fig. 261
bicolor Mart.
Leaf Globoid Brown No One-chambered TF Unidentified Fig. 262
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Grandez-Rios J.M. et al.

Table 3. Number and percentage of gall morphotypes in the different taxa 2011, Maia 2011, Araújo et al. 2012, Julião et al. 2017, Carvalho &
of galling insects recorded in the Allpahuayo-Mishana National Reserve and Mota 2018), a result also observed in the present study. Araújo et al.
Quistococha Regional Reserve, Iquitos, Peru.
(2012) suggested that the main explanation for the high insect gall
Gall-inducing taxa Gall morphotypes richness hosted by Fabaceae in the Amazon is the large number of
N % species in that family. According to Julião et al. (2014b), Fabaceae
Diptera (Cecidomyiidae) 58 22.14 stands out as the family with the greatest richness of Cecidomyiidae
Thysanoptera 2 0.76 galls in the Amazon. It is expected that the number of taxa in a host
plant family will be reflected in the number of galling associates, with
Lepidoptera 2 0.76
more species-rich families hosting a richer fauna (Gonçalves-Alvim
Hemiptera (Psyllidae) 1 0.38 & Fernandes 2001). Consistent with previous work, we showed that
Unidentified 199 75.95 the host genus Eschweilera (Lecythidaceae) was one of the richest in
Total 262 100 insect gall morphotypes. Julião et al. (2014b) found that 77% of galling
insect morphospecies sampled from Lecythidaceae are associated with
this single genus. With respect to the high gall diversity on T. guianesis,
Hemiptera (n = 1), and Lepidoptera (n = 1) (Table 2); white-sand C. utilis, M. elata, E. coriacea and S. parvifructa—the most important
dry forest was next, with Diptera (n = 15) and Lepidoptera (n = 1) as host species in the present study—there are no reports in the literature.
gall-inducing taxa. White-sand wet forest and palm swamp forest had Among all host plants, only C. utilis is endemic to the Amazonian
only Diptera (n = 19 and n = 4, respectively). Additionally, two gall- region. This plant hosts 14 gall-inducing species. Since galls result
inducing insects (Diptera) were found in two types of vegetation, one from species-specific interactions, the plant endemism also applies to
in palm swamp and white-sand dry forests on Virola pavonis (A. DC.) the inducers. So, these 14 cecidogenous species are endemic too.
A.C. Sm. (Myristicaceae), and other in terra firme and white-sand dry Leaves were the most frequently galled plant organ in our study. This
forests on Aspidosperma excelsum Benth. (Apocynaceae). pattern was observed in previous studies, including inventories carried out
The associated fauna was found in 66 gall morphotypes, with the in the Amazon forest (Mani 1964, Almada & Fernandes 2011, Maia 2011,
highest frequency of occurrence observed in the white-sand wet forest Silva et al. 2011, Araújo et al. 2012, Julião et al. 2017, Carvalho & Mota
(n = 23 morphotypes), followed by terrra firme forest (n = 18), white- 2018), and in other biomes (Rodrigues et al. 2014, Bergamini et al. 2017,
sand dry forest (n = 17), and palm swamp forest (n = 6). Marinho et al. 2023). According to Proença & Maia (2023), 81% of galls
occurred on leaves in the Amazon forest. This is probably because leaves
Discussion represent an abundant and renewable resource available to gall inducers
(Maia 2011). The globoid shape also was the most common one, together
We recorded 262 gall morphotypes across our sites in the Peruvian with the green color, glabrous surface and one-chambered galls. These
Amazon, a relatively high number compared to other areas with similar predominant characteristics are also consistent with previous studies in
vegetation. For example, Almada & Fernandes (2011) reported the the Amazon (Almada & Fernandes 2011, Maia 2011, Silva et al. 2011,
highest insect gall diversity (n = 309 insect gall morphogypes) in an Araújo et al. 2012, Julião et al. 2017, Carvalho & Mota 2018).
Amazonian forest site, FLONA-Saracá Taquera, Brazil. Other studies in Galling insects of the family Cecidomyiidae were the most important
the Brazilian Amazon recorded from 27 to 236 morphotypes (Almada & in the fauna studied. This result corroborates previous studies that pointed
Fernandes 2011, Maia 2011, Silva et al. 2011, Araújo et al. 2012, to the dominance of Cecidomyiidae in the Amazon forest (Proença & Maia,
Julião et al. 2017, Carvalho & Mota 2018). Thus, our study presents 2023). For example, Julião et al. (2014b), showed that Cecidomyiidae
the second highest diversity of galling insects recorded in Amazonian was responsible for 72% of gall morphospecies in the Amazon forest.
forest. We also found that the average number of gall morphotypes per This points to Cecidomyiidae as the family with the greatest diversity of
plant species was 3.5, which was high when compared to other locations galling insects in the Neotropical region (Gagné 1994).
in the Amazon—with average values of 1.2, 2.0, 1.7, 1.4 (Almada & Regarding the associated fauna, Thysanoptera, Hymenoptera, Acari,
Fernandes 2011, Maia 2011, Araújo et al. 2012, Julião et al. 2017). and Collembola were found as successors. Additionally, Curculionidae
Variations in the insect gall richness and number of gall morphotypes (Coleoptera) and Sciaridae (Diptera) were identified as cecidophages,
per plant species can be explained by differences in sampling method, which is the first documented record of this in Amazonian forest.
collecting effort and host plants sampled (Bergamini et al. 2017). Hymenoptera was the only insect order represented as a parasitoid.
Among the sampled vegetation types, the terra firme forest exhibited Among the Hymenoptera families, Eulophidae was the most frequent
the greatest richness of gall morphotypes and host species. This can be in our study. This pattern aligns with previous findings in the Brazilian
attributed to the exceptionally diverse and heterogeneous flora found Amazon, where Eulophidae has been reported as the predominant
in this forest type, which characterized by nutrient-rich soils and a parasitoid group (Carvalho & Mota 2018). In restinga areas of the
wide array of niches and ecological resources (Pitman et al. 2001). state of Rio de Janeiro (Atlantic forest), Eulophidae were also the most
This finding is consistent with results reported by Julião et al. (2014b), frequent and best represented parasitoids (Maia & Azevedo 2009), as
which demonstrated that terra firme forests have a greater number of well as in Cerrado (Maia & Silva 2021), showing that this pattern can
galling insect morphospecies per sampled area than other vegetation be widespread in South America.
types, such as várzea forest. The taxonomic resolution of gall-inducing insects in our study
Many inventories of gall diversity in the Amazon forest indicate that was low, which was also observed in previous studies in the Amazon.
Fabaceae is the most important host plant family (Almada & Fernandes Julião et al. (2014b) recorded 1,549 gall morphotypes in the Brazilian

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Biota Neotrop., 23(4): e20231568, 2023 25

Insect galls of the Peruvian Amazon

Amazon, and inducers of only 386 morphotypes were identified to Gerardo Flores Llampazo: contributed to data collection.
order or family level. In that study, gall-inducing insects were found to José Reyna Huaymacari: contributed to data collection.
belong to the orders Diptera (Cecidomyiidae), Hemiptera, Coleoptera, Maria Antonieta Pereira de Azevedo: resources, writing – review &
Lepidoptera, and Hymenoptera, while 24% of the inducers could not editing.
be identified to any possible level. Valéria Cid Maia: conceptualization, resources, methodology,
supervision, writing – review & editing.
Conclusions
Conflicts of Interest
This is the first systematic survey of insect galls undertaken in the
Peruvian Amazon. With this study, 262 gall morphotypes and 75 host plant The authors declare that they have no conflict of interest related to
species have been recorded. Fabaceae was the family with the highest the publication of this manuscript.
number of galled species. T. guianesis, C. utilis, M. elata, E. coriacea and
S. parvifructa were identified as the host plant species with the highest Data Availability
richness of gall-inducing insects. The leaf was the most galled plant organ,
and globoid shape was the most common morphology. Among the galler The datasets generated during and/or analyzed during the current
orders, Diptera was the most representative, with Cecidomyiidae being the study are available at: https://doi.org/10.48331/scielodata.025WUA.
standout family in terms of species richness. Hymenoptera was identified
as the only order of parasitoids, while successors were represented by four References
orders. Cecidophages were represented by both Coleoptera and Diptera ALMADA, E.D. & FERNANDES, G.W.A. 2011. Insetos indutores de galhas
orders. Finally, terra firme forest presented the largest richness of gall em florestas de terra firme e em reflorestamentos com espécies nativas na
morphotypes and host plant species. Amazônia Oriental, Pará, Brasil. Bol. Mus. Para. Emílio Goeldi. Cienc. Nat.
6(2):163–196.
Acknowledgments ARAÚJO, W.S.D. 2018. 30 years of research on insect galls in Brazil: a
scientometric review. Pap. Avulsos Zool. 58:1–11.
ARAÚJO, W.S.D., FERNANDES, G.W. & SANTOS, J.C. 2019. An overview
The first author wishes to thank the Programa Nacional de Becas y of inventories of gall-inducing insects in Brazil: looking for patterns and
Crédito Educativo (PRONABEC) for a scholarship for post-graduate identifying knowledge gaps. An. Acad. Bras. Ciênc. 91(1):e20180162.
studies in Brazil, and Dr. Gabriel Vargas Arana for the availability ARAÚJO, W.S.D., PORFÍRIO-JÚNIOR, É.D., JORGE, V.A. & ESPÍRITO-
of Laboratorio de química de productos naturales del Instituto de SANTO FILHO, K.D. 2012. Plantas hospedeiras e galhas entomógenas
em sub-bosques de florestas tropicais do Pará, Brasil. INSULA (41):59–72.
Investigaciones de la Amazonía Peruana (IIAP). We thank Julio Irarica
BERGAMINI, B.A.R., BERGAMINI, L.L., SANTOS, B.B.D. & ARAÚJO,
Pacaya, Aarón Panduro Bardales and Cecila Bardales Sandoval for their
W.S.D. 2017. Occurrence and characterization of insect galls in the Floresta
invaluable assistance during fieldwork. Dr. Adriano Cavalleri (UFRS) Nacional de Silvânia, Brazil. Pap. Avulsos Zool. 57(32):413–431.
by the Thysanoptera order identification, and Seth Kauppinen made CARDOSO, D., SÄRKINEN, T., ALEXANDER, S., AMORIM, A.M., BITTRICH,
comments on English usage. We also thank the Quistococha Tourist V., CELIS, M., DALY, D.C., FIASCHI, P., VIANA, P.L., STEHMANN, …
Complex, and the Peruvian Protected Areas Authority (SERNANP) STEVENS, W.D. 2017. Amazon plant diversity revealed by a taxonomically
verified species list. Proc. Natl. Acad. Sci. 114(40):10695–10700.
for providing research permits (RNAM N°11-2021-SERNANP-JEF)
for access to the forests the Allpahuayo-Mishana National Reserve. CARNEIRO, M.A.A., BRANCO, C.S.A., BRAGA, C.E.D., ALMADA, E.D.,
COSTA, M.B.M., MAIA, V.C. & FERNANDES, G.W. 2009. Are gall
E.N.H.C. also acknowledges support from a NERC Knowledge midge species (Diptera, Cecidomyiidae) host-plant specialists? Rev. Bras.
Exchange Fellowship (NE/V018760/1). Entomol. 53(3):365–378.
CARVALHO, A.N. & MOTA, J.S.D. 2018. Ocorrência e caracterização de galhas
Associate Editor entomógenas em um fragmento florestal em estágio de sucessão ecológica
na Amazônia. EntomoBrasilis 11(2):118–123.
Gustavo Graciolli DRAPER, F.C., HONORIO CORONADO, E.N., ROUCOUX, K.H.,
LAWSON, I.T., NIGEL, N.C., PAUL, P.V., PHILLIPS, O.L., TORRES
MONTENEGRO, L.A., VALDERRAMA SANDOVAL, E., MESONES, I.,
Author Contributions GARCÍA-VILLACORTA, R., ARÉVALO, F.R.R. & BAKER, T.R. 2018.
Peatland forests are the least diverse tree communities documented in
Julio Miguel Grandez-Rios: conceptualization, data curation, Amazonia, but contribute to high regional beta-diversity. Ecography
41(8):1256–1269.
research, methodology, resources, writing – original draft, writing –
ENCARNACIÓN, F. 1985. Introductión a la flora y vegetatión de la Amazonia
review & editing.
peruana : estado actual de los estudios, medio natural y ensayo de una clave
Walter Santos de Araújo: conceptualization, methodology, de determinación de las formaciones vegetales en la llanura amazónica.
supervision, writing – review & editing. Candollea 40(1):237–252.
Eurídice N. Honorio Coronado: contributed to data collection. ESPIRITO-SANTO, M.M. & FERNANDES, G.W. 2007. How many species
Timothy R. Bake: writing – review & editing, contributed to data of gall-inducing insects ere there on earth, and where are they? Annals of
the Entomological Society of America 100(2):95–99.
collection.
Rodolfo Vásquez Martínez: contributed to data collection. FORESTPLOTS.NET, BLUNDO, C., CARILLA, J., GRAU, R., MALIZIA, A.,
MALIZIA, L., OSINAGA-ACOSTA, O., BIRD, M., BRADFORD, M.,
Abel Monteagudo Mendoza: contributed to data collection. CATCHPOLE, … TRAN, H.D. 2021. Taking the pulse of Earth’s tropical forests
Roosevelt García-Villacorta: contributed to data collection. using networks of highly distributed plots. Biological Conservation 260:108849.

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Grandez-Rios J.M. et al.

GAGLIARDI-URRUTIA, G., GARCÍA DÁVILA, C., JARAMILLO- MAIA, V.C. & AZEVEDO, M.A.P. 2009. Micro-himenópteros associados com
MARTINEZ, A., ROJAS-PADILLA, O., RÍOS-ALVA, E., AGUILAR- galhas de Cecidomyiidae (Diptera) em Restingas do Estado do Rio de Janeiro
MANIHUARI, R., PÉREZ-PEÑA, P.E., TUPINAMBÁ, P., CASTROVIEJO- (Brasil). Biota Neotrop. 9(2):151–164.
FISHER, S., SIMÕES, P.I. & ESTIVALS, G. 2022. Anfibios de Loreto. 1 ed. MAIA, V.C. & SILVA, B.G. 2021. Insect galls of the Brazilian Cerrado:
Instituto de Investigación de la Amazonía Peruana, Peru. associated fauna. Biota Neotrop. 21(3):e20211202.
GAGNÉ, R.J. 1994. The gall midges of the Neotropical Region, 1st ed., Cornell MANI, M.S. 1964. Ecology of Plant Galls. 1nd ed, Junk, Haia. 434p.
Univ Press: Ithaca, New York. 325p.
MARENGO, J.A. 1998. Climatologia de la zona de Iquitos, Peru. In Geoecologia
GAGNÉ, R.J. & JASCHHOF, M. 2021. A Catalog of the Cecidomyiidae (Diptera) y desarrollo amazónico: estudio integrado en la zona de Iquitos, Peru (R.
of the World. 5th ed. Digital. 813 pp. Kalliola & S. Flores Paitan, eds) Annales Universitatis Turkuensis Ser A,
GIBSON, G. A. P., HUBER, J. T. & WOOLLEY, J. B. 1997. Annotated keys 114. University of Turku, Finland, p.35–57.
to the genera o Nearctic Chalcidoidea (Hymenoptera). NRC Research MARINHO, R.A., MAIA, V.C. & BARBOSA, M.R.D.V. 2023. Entomogenous
Press, Ottawa. galls and their associated fauna in deciduous dry forest and woodland
GONÇALVES-ALVIM, S.J. & FERNANDES, G.W. 2001. Biodiversity of vegetation remnants (Caatinga) in the Northern Depressão Sertaneja, Brazil.
galling insects: historical, community and habitat effects in four Neotropical Biota Neotrop. 23(1):e20221428.
savannas. Biodiversity and Conservation 10:79–98. MENDONÇA, M.D.S. 2007. Plant diversity and galling arthropod diversity
GRANDEZ-RIOS, J.M., GARCÍA-VILLACORTA, R., CUEVAS-REYES, searching for taxonomic patterns in an animal-plant interaction in the
P. & ARAÚJO, W.S. 2015. Insectos inductores de Agallas en América Neotropics. Bol. Soc. Argent. Bot. 42(3–4):347–357.
Latina: Ecología, Importancia y Nuevas Perspectivas. Rev. Biol. Neotrop. PHILLIPS, O., BAKER, T., FELDPAUSCH, T. & BRIENEN, R. 2009.
12(2):92–103. RAINFOR: field manual for plot establishment and remeasurement. The
ISAIAS, R.M.S., CARNEIRO, R.G.S., OLIVEIRA, D.C. & SANTOS, J.C. Royal Society, Leeds, UK.
2013. Illustrated and annotated checklist of Brazilian gall morphotypes. PITMAN, N.C.A., TERBORGH, J.W., SILMAN, M.R., NUN, P., CER, C.E.,
Neotrop. Entomol. 42:230–239. PALACIOS, W.A. & AULESTIA’, M. 2001. Dominance and distribution of
IUCN (INTERNATIONAL UNION FOR CONSERVATION OF NATURE). tree species in upper Amazonian terra firme forests. Ecology 82(8):2101–2117.
2023. The IUCN Red List of threatened species. Version 2023–1. http:// PROENÇA, B. & MAIA, V.C. 2023. Insect galls from Amazon rainforest areas
www. iucnredlist.org/ (last access on 14/12/2023). in Rondônia (Brazil). An. Acad. Bras. Cienc. 95(4):1–35.
JULIÃO, G.R., VENTICINQUE, E.M., FERNANDES, G.W. & PRICE, P.W. RODRIGUES, A.R., MAIA, V.C. & COURI, M.S. 2014. Insect galls of restinga
2014a. Unexpected high Diversity of Galling insects in the Amazonian upper areas of Ilha da Marambaia, Rio de Janeiro, Brazil. Rev. Bras. entomol.
canopy: the savanna out there. PLoS ONE 9(12):e114986. 58(2):173–197.
JULIÃO, G.R., ALMADA, E.D. & FERNANDES, G.W. 2014b. Galling ROUCOUX, K.H., LAWSON, I.T., JONES, T.D., BAKER, T.R.,
insects in the Pantanal wetland and Amazonian rainforest. In Neotropical CORONADO, E.N.H., GOSLING, W.D. & LÄHTEENOJA, O. 2013.
Insect Galls (G. W. Fernandes & J. C. Santos, eds) Springer Netherlands, Vegetation development in an Amazonian peatland. Palaeogeography,
Dordrecht, p.377–403. Palaeoclimatology, Palaeoecology 374:242–255.
JULIÃO, G.R., ALMADA, E.D., COSTA, F.R.C., CARNEIRO, M.A.A. & SALO, M. & PYHÄLÄ, A. 2007. Exploring the gap between conservation
FERNANDES, G.W. 2017. Understory host plant and insect gall diversity science and protected area establishment in the Allpahuayo-Mishana
changes across topographic habitats differing in nutrient and water stress in National Reserve (Peruvian Amazonia). Envir. Conserv. 34(1):23–32.
the Brazilian Amazon rainforest. Acta Amaz. 47(3):237–246.
SILVA, P.S.D., ALMEIDA-SANTOS, B., TABARELLI, M. & ALMEIDA-
LEÓN, B., PITMAN, N. & ROQUE, J. 2006. Introducción a las plantas CORTEZ, J.S. 2011. Occurrence of gall complexes along a topographic
endémicas del Perú. Rev. Peru. Biol. 13(2):9s–22s. gradient in an undisturbed lowland forest of central Amazonia. Revista
LOPEZ-GONZALEZ, G., LEWIS, S.L., BURKITT, M. & PHILLIPS, O.L. Brasileira de Biociências 9(2):133–138.
2011. ForestPlots.net: a web application and research tool to manage STONE, G.N. & SCHÖNROGGE, K. 2003. The adaptive significance of insect
and analyse tropical forest plot data. Journal of Vegetation Science gall morphology. Trends in Ecology & Evolution 18(10):512–522.
22(4):610–613.
YUKAWA, J., TOKUDA, M., UECHI, N. & SATO, S. 2001. Species richness of
LUZ, F.A. & MENDONÇA JÚNIOR, M.S. 2019. Guilds in Insect Galls: Who galling arthropods in Manaus, Amazon and the surroundings of the Iguassu
is Who. Florida Entomol. 102(1):207. Falls. Esakia 41:11–15.
MAIA, V.C. 2011. Characterization of insect galls, gall makers, and associated
fauna of Platô Bacaba (Porto de Trombetas, Pará, Brazil). Biota Neotrop.
11(4):37–53. Received: 22/09/2023
MAIA, V.C. 2022. Review of the inquilinous fauna associated with insect galls Accepted: 28/12/2023
in Brazilian restingas. Brazilian J. Biol. 821–15. Published online: 26/01/2024

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