Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

The insects of the inflorescences of neotropical palms (Arecaceae):

diversity and coevolution

Cesar Delgado1*; https://orcid.org/0000-0002-4961-7736

Joel Vasquez2; https://orcid.org/0000-0002-0584-7310

Kember Mejía1; https://orcid.org/0000-0002-2634-7227

Guy Couturier3; http://orcid.org/0009-0002-1966-060X

1. Departamento de Diversidad Biológica Amazónica Terrestre, Instituto de Investigaciones de la Amazonia Peruana,

Avenida José Abelardo Quiñones km 2.5, San Juan, Iquitos, Perú; cdelgado@iiap.gob.pe (*Correspondencia), kmejía@

iiap.gob.pe

2. Facultad de Ciencias Forestales, Universidad Nacional de la Amazonía Peruana, Carretera Zungarococha, Puerto

Almendra 16000, San Juan, Loreto, Perú; joel.vasquez@unapiquitos.edu.pe

3. Département Systématique et Evolution, Muséum national d’Histoire naturelle, rue Cuvier, 75231 Paris Cedex 05,

France; guy.couturier4@wanadoo.fr

Received 04-VII-2025. Corrected 17-I-2026. Accepted 09-IV-2026.

ABSTRACT

Introduction: Palms hold a significant position in the Amazonian flora. This study examines the insects associ-

ated with the inflorescences of various genera and species of neotropical palms, with a focus on those of local or

national economic significance in the countries studied (Brazil, Ecuador, Peru).

Objective: To characterize the entomological community and analyze the diversity of insects in several species

and genera of neotropical palm trees, and to demonstrate the coevolutionary relationships between insects and

palm trees.

Methods: The study was conducted between 1990 and 2020, in the Brazilian, Ecuadorian, and Peruvian Amazon,

in the Atlantic Forest of Brazil, and in the Peruvian Andes. Several thousand insects were collected in plastic bags

or with an entomological net on the inflorescences of selected palm trees and anesthetized with insecticides. The

insects were classified into families. A significant part was prepared on entomological pins, referenced, and sent

to specialists for identification. Species new to science were described whenever possible.

Results: Consistent with previous studies, it is confirmed that the Nitidulidae (Mystropini) and Curculionidae

(Derelomini) families are specialists in palm inflorescences. Additionally, among hemipterans, the families

Miridae (Phylini) and Thaumastocoridae (Discocoris), as well as Staphylinidae beetles, exhibit a high degree of

specialization. In contrast, the Dynastidae (Cyclocephala genus) and Apidae (Trigoninae), frequently cited as

palm pollinators, are not specialized and should be considered as opportunistic.

Conclusion: The specific-level analysis presented here shows that a close coevolutionary relationship exists

between certain insect species and specific genera or species of palms.

Key words: Amazonia; Atlantic Forest; Andes; pollinators.

https://doi.org/10.15517/7zr3v322

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

INTRODUCTION

Palms hold a significant position in the

Amazonian flora and have been studied by var-

ious authors (Bernal & Galeano, 2006; Drans-

field & Beentje, 1996; Henderson et al., 1995;

Kahn & de Granville, 1992; Kahn & Mejia,

1988; Kahn & Mejía, 1990; Lima et al., 1986;

Uhl & Moore, 1977). It is estimated that there

are currently 39 genera and 180 described spe-

cies native to the Amazon, with the Ceroxylon

genus, found beyond the western boundary

of the Amazon (Kahn & de Granville, 1992),

along with the palm Attalea funifera from the

Brazilian Atlantic Forest (Voecks, 2002). Since

the synthesis conducted by Henderson (1986),

insects associated with palm inflorescences

have been the subject of numerous publications

due to their relevance to pollination. Barfod et

al. (2011) provide a critical review of studies

published over the past 25 years, particularly

mentioning the negative role of certain insects

in the reproductive success of palms. Addition-

ally, Howard et al. (2001) as, more recently,

Henderson (2024) published a synthesis of our

knowledge on palm-associated insects.

The objective of this work is to character-

ize the populations of different insect species

in relation to the various species and genera

of the most common neotropical palms to

demostrate an evident coevolution. Insects that

clearly attack inflorescences before they open,

such as Foveolus spp. (Curculionidae), Dynamis

spp. (Dryophthoridae), and Hemiphileurus sp.

(Dynastidae), are not covered in this study.

These results were obtained from over

20 years of observations by the authors in the

Brazilian, Ecuadorian, and Peruvian Amazon.

The palm species selected for the monitoring

were those used by humans for food and medi-

cine, as well as for various domestic purposes

(Balick, 1984; Balslev et al., 2008; Brañas &

Horna, 2011; Brokamp et al., 2011; Collazos &

Mejía, 1988; Didonet & Ferraz, 2014; Graefe

et al., 2013; Instituto de Investigaciones de

la Amazonía Peruana [IIAP], 2005; Lima et

al., 1986; Kahn, 1988; Mejía & Kahn, 1996;

Moussa & Kahn, 1997; Moussa et al., 1996;

Paniagua-Zambrana et al., 2017). Some of these

are of great economic importance in local and

national markets (Clement & Mora-Urpí, 1987;

Delgado et al., 2007; Mejía, 1992; Barboza da

RESUMEN

Los insectos de las inflorescencias de palmas neotropicales (Arecaceae): diversidad y coevolución

Introducción: Las palmas ocupan un lugar destacado en la flora amazónica. Este estudio examina los insectos

asociados con las inflorescencias de varios géneros y especies de palmas neotropicales, centrándose en aquellos

de importancia económica local o nacional.

Objetivo: Caracterizar la comunidad entomológica y analizar la diversidad de insectos en varias especies y géne-

ros de palmeras neotropicales, y demostrar las relaciones coevolutivas entre insectos y palmeras.

Métodos: El estudio fue conducido entre los años 1990 y 2020, en la Amazonía brasileña, ecuatoriana y peruana,

en la Mata Atlántica de Brasil y en los Andes peruanos. Varios miles de insectos fueron recolectados en bolsas de

plástico o con red entomológica en las inflorescencias de las palmeras seleccionadas y anestesiados con insectici-

das. Los insectos fueron separados por familias. Una parte significativa fue preparada en alfileres entomológicos,

referenciada y enviada a especialistas para su identificación. Las especies nuevas para la ciencia fueron descritas,

cada vez que fue posible.

Resultados: Concordante con estudios previos, las familias Nitidulidae (Mystropini) y Curculionidae (Derelomini)

son especialistas en inflorescencias de palmeras. Además, entre los hemípteros, las familias Miridae (Phylini) y

Thaumastocoridae (Discocoris), así como los escarabajos Staphylinidae, exhiben un alto grado de especialización.

En contraste, los Dynastidae (género Cyclocephala) y Apidae (Trigoninae), frecuentemente citados como polini-

zadores de palmeras, no están especializados y deben considerarse como oportunistas.

Conclusión: El análisis a nivel específico presentado aquí muestra que existe una estrecha relación coevolutiva

entre ciertas especies de insectos y los géneros o especies de palmeras estudiados.

Palabras clave: Amazonía; Mata Atlántica; Andes; polinizadores.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

Silva et al., 2021). These results, many of which

had not been previously published, are comple-

mented by those of other authors published in

the last 30 years.

MATERIALS AND METHODS

Study areas: The study was primarily con-

ducted in the Amazon region: in Brazil, in the

states of Pará, at Empresa Brasileira Agropecu-

aria (Embrapa) (Belém, 1°08’34’’ S & 48°43’16’’

W) and Amazonas (Manaus, Rio Urubu, 02°06’

S & 60°02’ W, Rio Negro community Terra

Preta, 02°30’40’’ S & 60°52’34’’ W); in Ecuador,

in the provinces of Sucumbíos (Shushufindi,

0°03’15’’ S & 77°06’58’’ W), Morona Santiago

(San Rafael, l0°09’82’’ S & 77°16’71’’ W), Santo

Domingo (Quinindé, Palmeras de Los Andes,

00°13’11’’ S & 79°06’58’’ W), and Cotopaxi

(Otonga Reserve, 02°02’08’’ S & 78°51’04’’ W);

in Peru, in the departments of Loreto (Iquitos,

El Dorado research station of the National Agri-

cultural Research Institute (INIA), 03°57’16’’ S

& 73°24’41’’ W, Jenaro Herrera, Instituto de

Investigaciones de la Amazonía Peruana (IIAP)

research station, 04°54’14’’ S & 73°40’50’’ W

(IIAP), Iquitos-Nauta road 04°32’ S & 73°33’ W,

Allpahuayo Reserve, 03°97’89’’ S & 73°42’19’’

W, Yanapuma site, 04°23’44’’ S & 73°34’23’’ W,

Nauta city (la Circular, 04°39’ S & 73°35’ W),

Maniti(Santa María, 03°36’’ S & 72°57’ W) and

Rio Yarapa near Puerto Miguel 73°23’13’’ W

& 04°29’21’’ S, in the department of Madre de

Dios, Puerto Maldonado 12°50’ S & 59°20’ W,

and in the department of San Martín, Uchiza

(Palmas del Espino plantation 08°27’33’’ S &

76°27’48’’ W). Outside the Amazon, sampling

was conducted in the Atlantic Forest, in the

state of Bahia (Brazil: Ilheus-Itabuna 14°47’ S &

39°21’ W), and in the Peruvian Andes, depart-

ment of Amazonas (Ocol, 2 370 m altitude

06°15’48’’S & 77°34’41’’ W).

Collection methods and preservation

of entomological material: Inflorescences in

anthesis were collected in transparent plastic

bags, whose dimensions were adapted to those

of the inflorescences. In some cases, only part

of the inflorescence was collected due to its

large size (e.g., Mauritia flexuosa), or directly

with an entomological net. This allowed for the

collection of most insects present in the target-

ed part of the plant. Sometimes it was necessary

to climb the stipe to reach the inflorescence.

Insects were quickly anesthetized with insec-

ticidal spray and placed in the shade to avoid

tissue swelling. In the laboratory or a suitable

location, the insects were separated from the

plant material present in the sample (anthers

and other plant debris) and placed in 70 % alco-

hol with proper references. Later, insects were

sorted by families, and representative samples

were sent to specialists for specific identifica-

tion. New species for science were described.

It should be noted that in the case of the Cur-

culionidae Derelomini, which are difficult to

identify, not all specimens were identified at the

species level; some even require a revision of

several genera (O’Brien, pers. comm., 2014). In

these cases, they were labelled with a letter for

identification purposes.

Identifications and material deposits:

The identified species are housed in the col-

lections of the following museums and labora-

tories: Museum of Entomology at the National

Agrarian University La Molina (Peru), Peru-

vian Amazon Research Institute, Iquitos (Peru),

National Museum of Natural History (Paris),

entomological collection of the Empresa

Brasileira de Pesquisa Agropecuaria (Embra-

pa, Belém, Brazil), entomological collection of

the National Institute for Amazonian Research

(INPA, Manaus, Brazil), Museum of Natural

History of Rio de Janeiro (Brazil), the collec-

tion was destroyed due to the fire of 2 Septem-

ber 2018, private collection of Roberto Pace

(Verona, Italy), private collection of Charles

O’Brien (Green Valley, Arizona, United States

of America) now in the collection of the Ari-

zona State University, (Tempe, United States of

America), Museum of Natural History of Saint

Petersburg (Russia). Some genus and species

names of palms have been modified during this

work. The names mentioned are those from

the time of collection. Synonyms are provided

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

in parentheses. The palms were identified by F.

Kahn and K. Mejia, botanists and specialists in

neotropical palms.

RESULTS

The results are organized by palm genera

and species, followed by notable insects. Given

that the taxonomy of palms has been modified,

we found it preferable to maintain the identifi-

cations as they appear on the collection labels,

indicating synonyms here. The Astrocaryum

genus has been extensively reviewed by Kahn

(2008) and Kahn and Millán (1992). The Atta-

lea genus has been reviewed by Zona (2002)

and Pintaud (2008). Balick (1986) reviewed the

Oenocarpus or Jessenia complex.

The palms

Attalea spp. (= Scheelea, Orbignya, Maxi-

miliana): The genus was reviewed by Pintaud

(2008) and Zona (2002). Two species were

studied: A. funifera Mart. Ex Spreng. in the

Atlantic Forest (Brazil) and Attalea plowmanii

(Glassman) in the Allpahuayo Reserve area

(Iquitos, Peru).

Astrocaryum subgender, Monogynanthus

and A. subg. Pleiogynanthus: The genus was

reviewed by Kahn and Millán (1992), and we

sampled the following species:

• Subgenus Monogynanthus: Astrocaryum

gratum Kahn & Millan, Astrocaryum java-

rense Trail ex Drude, Mart., Astrocaryum

macrocalyx and Astrocaryum urostachys

Burrett.

• Subgenus Pleiogynanthus: Astrocaryum

aculeatum Meyer, Astrocaryum jauari

Martius, Astrocaryum chambira Burrett,

Astrocaryum vulgare Martius. Consiglio

and Bourne (2001) reported about a sin-

gle inflorescence of A. vulgare in Guyana,

with nearly 30 000 Nitidulidae (possibly

Mystropini).

Padilha de Oliveira et al. (2003) studied

the pollination biology of A. vulgare in Brazil

(Belém), and T. Peyret (nov. 2000, unpublished)

studied A. urostachys in Ecuador. Several spe-

cies of the genus Celetes develop in young fruits:

in a count on A. javarense in Iquitos and A. gra-

tum in Puerto Maldonado, we found between

95 and 100 % of fruits parasitized (Table 1).

Table 1

Parasitism of the female flowers of Astrocaryum javarense and Astrocaryum gratum by the genus Celetes sp. (Derelomini) in

three locations in Peru.

Palm Locality Total number

of female flowers

Number of

parasitized flowers

Formed

fruits

Celetes

spp.

A. javarense (Loreto, Tamshiyacu) Inflorescence

A 342 322 20 sp. 1

B 301 301 0 sp. 1

C 367 367 0 sp. 1

A. javarense (Loreto, Santa Cecilia) Inflorescence

A 377 303 + 74 (*) 0sp. 1

B 111 98 + 13 (*) 0sp. 1

A. gratum (Madre de Dios, Puerto Maldonado Inflorescence

A 414 414 0 sp. 1

sp. 2

sp. 3

(*) second number: undeveloped fruits, but with doubts about parasitism.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

Bactris gasipaes Kunth: From an econom-

ic perspective, B. gasipaes is the most studied

species (Beach, 1984; Clement & Mora-Urpí,

1987; Essig, 1971; Graefe et al., 2013; Mora-

Urpí & Mexzon, 1996; Mora-Urpí & Solís, 1980;

Oviedo et al., 2020). We sampled it in several

locations in Brazil, Ecuador, and Peru.

B. gasipaes is characterized by the presence

of Anomalocornis couturieri (Miridae) (see Het-

eroptera below), Mystrops bactrii (Nitidulidae),

Phyllotrox spp., and several other species of

Derelomini (Listabarth, 1996). Mora-Urpí and

Solís (1980) reported Andranthobius (= Derelo-

mus) palmarum.

Ceroxylon quindiuense (H. Karst.) H.

Wendl., an Andean palm, has been reviewed

by Galeano et al. (2008), Sanín and Galeano

(2011), and Sanín et al. (2013). The entomo-

logical fauna of C. quindiuense presents signifi-

cant particularities, including the presence of

the Derelomini Terioltes mejiai (Perrin & Cou-

turier, 2024), six species of Mystropini (Kire-

jtshuk & Couturier, 2009) (Table 2), and the

Cyclocephalinae Ancognatha vulgaris Arrow.

In a sampled male inflorescence, we estimated

a total of around 250 000 insects. Carreño-Bar-

rera et al. (2019) and Carreño-Barrera (2020)

studied the entomological fauna of Ceroxylon

parvifrons (Engel.) H. Wendl., Ceroxylon ven-

tricosum Burret, and Ceroxylon vogelianum

(Engel.) H. Wendl. in Colombia. They reported

A. vulgaris and seven unidentified species of

Staphylinidae Aloeocharinae. Regarding the

Nitidulidae Mystropini, they found the same

species as we did on C. quindiuense, confirm-

ing the specialization of these insects in the

Ceroxylon genus. Balhara et al. (2013) studied

the floral structure of Ceroxylon ceriferum in

Venezuela.

• In Oenocarpus bataua in Ecuador and Perú

(Pace, 2010).

• Amazoncharis: 3 unidentified species in

Phytelephas seemanni in Colombia (Bernal

& Ervik, 1996).

• In Orbignya polysticha (Attalea) (Pace,

2010).

Elaeis oleifera (Kunth) Cortés and Elaeis

guineensis Jacq.: E. guineensis, the African

oil palm, was included in this study due to its

close relationship with E. oleifera. Three spe-

cies of Derelomini native to Africa Elaeidobius

kamerunicus Faust, Elaeidobius singularis Faust,

and Elaeidobius plagiatus F. were intentionally

introduced to this palm to improve pollination

and, thus, productivity (Genty et al., 1986; Auf-

fray et al., 2017). A fourth species, Elaeidobius

subvittatus Faust, like the other three, is found

on E. guineensis. It was likely introduced much

earlier, possibly during the slave trade. The

species was described by Bondar as Elaeisae

in 1942 (O’Brien & Woodruff, 1986). Haran

et al. (2023) also recently reviewed the genus

and described two new species of African

Elaeidobius. E. oleifera has its own fauna and

does not host any Elaeidobius species, with

the Derelomini Grasidius hybridus, Grasidius

sp., Couturierius carinifrons, and Couturierius

constrictirostris being strictly dependent on E.

oleifera (O’Brien et al., 2004; Beaudoin-Ollivier

et al., 2017).

Regarding the hybrid E. guineensis × E. ole-

ifera, Meléndez and Ponce (2016) demonstrated

that the first-generation hybrid attracts Derelo-

mini from both species. However, the backcross

E. guineensis × E. oleifera × E. guineensis loses its

appeal to Derelomini associated with E. oleifera

(Couturier et al., 1996) (Table 3).

Euterpe oleracea Engel. and Euterpe prec-

atoria Mart.: Campbell et al. (2018) and Bezer-

ra et al. (2020) show that 90 % of pollination is

carried out by native bees (Table 4).

Different species of Cyclocephalini known

in palm inflorescences:

• A. vulgaris arrow on C. quindiuense.

• Cyclocephala amazona L. on A. macrocalyx,

B. gasipaes, O. bataua, M. flexuosa, Phyte-

lephas tenuicaulis, and E. oleifera.

• Cyclocephala boulardii Dechambre on Bac-

tris hirta (Küchmeister et al., 1998).

• Cyclocephala collaris Burmeister on E.

oleifera.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

Table 2

Presence of different species of beetles: Curculionidae, Nitidulidae, and Staphylinidae in the inflorescences of palm trees.

Astrocaryum

subgenera

Monogynanthus

Astrocaryum

subgenera

Pleiogynanthus

Attalea

spp.

Bactris

gasipaes

Bactris maraja,

B. monticola

Ceroxylon

quindiuense

Elaeis

oleifera

Elaeis

guineensis

Euterpe oleracea,

E. precatoria

Mauritia flexuosa,

M. carana

Oenocarpus

spp.

Phytelephas

tenuicaulis

Socratea sp.

Iriartea sp.

Syagrus

spp.

Curculionidae

Anchylorhynchus spp.0 0 0 0 0 0 0 0 0 0 ++ 0 0 ++

Celetes spp. AA +++ 0

Celetes spp. BB +++ 0

Couturierius spp. 0 0 0 0 0 0 +++ 0 0 0 0 0 0 0

Grasidius spp. 0 0 0 0 0 0 +++ 0 0 0 0 0 0 0

Elaeidobius spp.0 0 0 0 0 0 0 +++ 0 0 0 0 0 0

Terires sp.0

Terires minusculus ++ ++ 0 0 0 0 0 0 0 0 0 0 0 0

Phyllotrox spp.+ 0 0

Nitidulidae

Anthepurops depressa 0 0 0 0 0 0 0 0 0 0 0 +++ 0 0

Anthocorcina subcalva 0 0 0 0 0 0 0 0 0 0 0 0 +++ 0

Mystrops astrocaryi +++ 0 0 0 0 0 0 + 0 0 0 0 0 0

M. atrata 0 0 0 0 0 0 0 0 0 0 0 +++ 0 0

M. bactrii 0 0 0 +++ 0 0 0 0 0 0 0 0 0 0

M. beserrai 0 0 0 0 0 0 + 0 0 0 +++ 0 0 0

M. costaricensis 0 0 0 0 0 0 +++ +++ 0 0 0 0 0 0

M. dalmasi 0 0 0 0 0 0 0 0 0 +++ 0 0 0 0

M. debilis (1) 0 0 +++ 0 0 0 0 0 0 0 0 0 0 0

M. delgadoi 0 0 0 0 0 +++ 0 0 0 0 0 0 0 0

M. discoidea 0 0 0 0 0 0 0 0 0 0 0 0 0 0

M. gigas 0 0 0 0 0 ++ 0 0 0 0 0 0 0 0

M. hisamatsui 0 0 0 0 0 ++ 0 0 0 0 0 0 0 0

M. kahni +++ 0

M. komissari 0 0 0 0 ++ 0 0 0 0 0 0 0 0 0

M. lobanovi 0 0 0 0 0 0 0 0 0 0 0 0 +++ 0

M. neli 0 0 0 0 0 +++ 0 0 0 0 0 0 0 0

M. pectoralis 0+++ 0 0 0 0 0 0 0 0 0 0 0 0

M. pulchra 0 0 0 0 0 +++ 0 0 0 0 0 0 0 0

M. rotundula 0 0 0 0 0 +++ 0 0 0 0 0 0 0 0

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

• Cyclocephala couturieri Dechambre on

Astrocaryum sp. aff. murumuru.

• Cyclocephala decolorata Herbst on A. uros-

tachys, O. bataua, and M. flexuosa.

• Cyclocephala distincta Burmeister on

B. gasipaes, E. oleifera, and Oenocarpus

mapora.

• Cyclocephala forsteri Endrödi on Acroco-

mia aculeata (Scariot et al., 1991).

• Cyclocephala goetzi Endrödi 1966 on A.

urostachys.

• Cyclocephala guianae Endrödi 1969 on M.

flexuosa.

• Cyclocephala lunulata Burmeister on A. sp.

aff. murumuru.

• Cyclocephala marginalis Kirsch on A. uros-

tachys, O. bataua, and Attalea microcarpa

(Küchmeister et al., 1998).

• Cyclocephala rondoniana Ratcliffe on

Astrocaryum sp. and Attalea attaleoides

(Küchmeister et al., 1998).

• Cyclocephala stictica Burmeister on A.

urostachys.

• Cyclocephala undata Olivier on B. hirta

(Küchmeister et al., 1998).

• Cyclocephala variipenis Miki on A.

urostachys.

• Mimeoma signatoides Höhne on A. urosta-

chys, O. bataua, and O. mapora.

Lepidocaryum tenue Mart., a small under-

story palm with very inconspicuous inflo-

rescences, does not host any specialized and

generalist fauna in the different places where it

has been monitored.

M. flexuosa L. f. and Mauritia carana

Wallace: M. flexuosa is a dioecious palm abun-

dant in the Amazonian floodplains (aguajales).

We sampled M. flexuosa in Brazil, Ecuador, and

Peru. Exploited for its fruits (Delgado et al.,

2007), the “aguaje” hosts a very specific fauna,

primarily the Miridae: Phylini: Alvarengamiris

alvarengai, Alvarengamiris kemberi, Alvaren-

gamiris margaridae (Peyret et al., 2005), the

Staphylinidae Delgadobius amazonensis (Chani-

Posse & Couturier, 2012), the Thaumastocori-

dae Discocoris drakei (Couturier et al., 1998),

Astrocaryum

subgenera

Monogynanthus

Astrocaryum

subgenera

Pleiogynanthus

Attalea

spp.

Bactris

gasipaes

Bactris maraja,

B. monticola

Ceroxylon

quindiuense

Elaeis

oleifera

Elaeis

guineensis

Euterpe oleracea,

E. precatoria

Mauritia flexuosa,

M. carana

Oenocarpus

spp.

Phytelephas

tenuicaulis

Socratea sp.

Iriartea sp.

Syagrus

spp.

M. squamae +++ 0 0 0 0 0 0 0 0 0 0 0 0 0

M. vasquezi 0 0 0 0 0 0 0 0 0 0 +++ 0 0 0

Platychorodes adentatus 0 0 0 0 0 0 0 0 0 0 0 +++ 0 0

Staphylinidae

Delgadobius amazonensis 0 0 0 0 0 0 0 0 + +++ + 0 0 0

Amazoncharis spp. - - - - - 0 - - 0 - - +++(2) - -

Plesiomalota palmarum - - ++(3) - - - - - 0 - - -

Arecaceopora spp.0 0 0 0 0 0 0 0 0 0 0 +++ 0 0

Oussipaliaglossa couturieri 0 0 +++ 0 0 0 0 0 0 0 0 0 0 0

Atheta chunitayensis 0 ++

Atheta couturieri - - - - - - - - 0 - - ++ - -

Phoinixiusa ecuadorensis - - - - - - - - 0 - ++(1) - - -

Feluva pichinchaensis - - ++(3) - - -- - - - - - - - -

0 Absence, + Less than 10 in an inflorescence, they are considered erratic, ++ Between 10 and 100 in an inflorescence, +++ More than 100 in an inflorescence, - Not confirmed.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

Table 3

Number of individuals collected per inflorescence of different pollinating Derelomini species in E. guineensis, E. oleifera, the

hybrid E. guineensis × E. oleifera, and the backcross E. guineensis × (E. oleifera × E. guineensis) at the ‘Urubu River’ plantation,

Embrapa, Manaus, Brazil, October 1994.

Male inflorescence of Elaeidobius

kamerunicus

Elaeidobius

Subvittatus

Grasidius

hybridus

Couturierius carinifrons

y Couturierius constrictirostris

Elaeis guineensis 515 17

Elaeis oleifera 1 465 194

Elaeis oleifera 280 29

Elaeis oleifera 180 385

Elaeis oleifera. (end of anthesis) 7 1

Elaeis oleifera 648 4

Elaeis oleifera 937 319

E. guineensis x E. oleífera 1 567 2 250 3

E. guineensis x E. oleífera 3 521 1 940 20

E. guineensis x E. oleifera x guineensis 234

E. guineensis x E. oleifera x guineensis 8 661

Table 4

Presence of different species of Apidae in the sampled palm trees.

Genus, species Host plant Country, province, department or state, locality

(Collectors, Authors)

Apis mellifera LMauritia flexuosa Perú, Loreto, Santa Cecilia, (KM, 1991)

Melipona ebúrnea Friese Mauritia flexuosa Perú, Loreto, Santa Cecilia, (KM, 1991)

Melipona fasciata Cockerell Phytelephas seemanni Ecuador

Partamonia epiphytophila Pedro & Camargo Mauritia flexuosa Perú, Loreto, Santa Cecilia (KM, 1991)

Partamona vicina Camargo Euterpe oleracea Brasil, Para, Belem Abaetetuba (GC, FK, 1996)

Ptilotrigona lurida Smith Astrocaryum acaule (Küchmeister et al., 1998)

Tetragona clavipes Fab. Mauritia flexuosa Perú, Loreto, Santa Cecilia (KM, 1991)

Trigona amalthea Olivier Mauritia flexuosa Perú, Loreto, Iquitos (KM, 1991, 1992)

Trigona amalthea Olivier Phytelephas seemanni Ecuador Bernal & Ervik

Trigona amazonensis Ducke Ceroxylon quindiuense Perú, Amazonas, Ocol (GC, CD, KM, 2007)

Trigona amazonensis Ducke Attalea funifera Brasil, Para, Ilheus (GC, JD, 1993)

Trigona amazonensis Ducke Maximiliana maripa (Oenocarpus)Brasil, Amazonas, Manaus (GC, 1997)

Trigona cf. brauneri Cockerell Astrocaryum gynacanthum Brasil, Amazonas, Manaus, Küchmeister et al., 1998)

Trigona cf. brauneri Cockerell Attalea attaleoides Brasil, Amazonas, Manaus Küchmeister et al., 1998)

T. chanchamayoensis Schwarz Phytelephas seemanni Ecuador (Bernal & Ervik, 1996)

Trigona dellatorreana Friese Euterpe oleracea Brasil, Para, Belem Abaetetuba (GC, FK, 1996)

Trigona dellatorreana Friese Mauritia flexuosa Perú, Loreto, Santa Cecilia (KM, 1991)

T. ferricaudata Cockerell Phytelephas seemanni Ecuador (Bernal & Ervik,1996)

Trigona fuscipennis Friese Unidentified palm Brasil, Para, Manacaparamirim (GC, FK 1996)

Trigona hyalinata Lepeletier Elaeis oleífera Brasil, Bahia, Ilheus (GC, JD, 1993)

T. hyalinata Lepeletier Phytelephas seemanni Ecuador (Bernal & Ervik, 1996)

T. nigerrima Cresson, Phytelephas seemanni Ecuador (Bernal & Ervik, 1996)

T. schultesii Friese Phytelephas seemanni Ecuador (Bernal & Ervik, 1996)

T. spinipes Fab., Phytelephas seemanni Ecuador (Bernal & Ervik, 1996)

T. williana Friese, Phytelephas seemanni Ecuador (Bernal & Ervik, 1996)

Trigona williana Friese Attalea microcarpa (Küchmeister et al., 1998)

T. fulviventris Guérin-Menneville Attalea microcarpa (Küchmeister et al., 1998)

Collectores: GC: G. Couturier, MJC: M. J. Cravo, JD: J. Delabie, FK: F. Kahn, KM: K. Mejía.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

and the Nitidulidae Mystrops dalmasi. All of

these species reproduce on the male inflo-

rescence where they are very abundant. The

Derelomini have not been identified by us, but

Núñez-Avellaneda and Carreño-Barrera (2013)

studied them in greater detail.

Oenocarpus (= Jessenia): The species

considered are O. bataua Mart., Oenocarpus

bacaba Mart., Oenocarpus balickii Kahn, Oeno-

carpus distichus Mart., O. mapora H. Karst.,

and Oenocarpus multicaulis Spruce. All the

sampled species exhibit the same characteris-

tics with a highly specialized fauna (Tables 2,

Table 5). The characteristic insects include the

Thaumastocoridae D. drakei Slater & Ashlock,

present in all the species considered, as well

as various species of Anchylorhynchus (Anchy-

lorhynchus bicarinatus O’Brien on O. mapora

in Belém, Brazil, Embrapa). The latter are also

present in different species of Syagrus studied

by other authors (see “Curculionidae” below).

Núñez-Avellaneda et al. (2015) on three species

of Oenocarpus, Núñez-Avellaneda and Rojas-

Robles (2008) on O. bataua, Valente & Madeiros

(2013), Silberbauer-Gottsberger et al. (2013)

and de Melo-Valente and Souza de Medeiros

(2013), report various species of Anchylorhyn-

chus in several species of Syagrus (see below).

We found different species of Anchylorhynchus

in all the sampled species of Oenocarpus.

Phytelephas seemannii O.F. Cook, P. te n u -

icaulis (Barfod) A. J. Hend.: P. seemannii has

been studied in Ecuador by Bernal & Ervik

(1996), Barfod et al. (1987), Ervik et al. (1999),

while P. tenuicaulis was studied by us in Peru. P.

tenuicaulis is a multi-stemmed dioecious palm

of the lowlands (Costa et al., 2009). The male

inflorescences exhibit a great diversity of insects

(Table 2, Table 5), most of which develop on

them. In P. tenuicaulis, we found Mystropini

with three species, (Kirejtshuk & Couturier,

2010), an unidentified species of Derelomini,

not found on other palms, as well as Dro-

sophilidae larvae and Ptiliidae adults during

inflorescence decay. It is noteworthy that we

did not observe any insects on the female

Table 5

Presence of different species of heteropterans in the inflorescences of palm trees.

Astrocaryum subgénero

Monogynanthus

Astrocaryum Subgénero

Pleiogynanthus

Attalea

spp.

Bactris

gasipaes

Bactris maraja

B. monticola

Ceroxylon

quindiuense

Elaeis

oleifera

Elaeis

guineensis

Mauritia flexuosa,

M. carana

Mauritiella sp.

Oenocarpus spp. Phytelephas

tenuicaulis

Socratea

spp.

Syagrus

spp.

Miridae Fulvini

Anomalocornis spp. +++ +++ 0 +++ - 0 0 0 0 0 0 0 0

Alvarengamiris spp. 0 0 0 0 0 0 0 0 +++ 0 0 0 0

Fulvius sp. ++ 0 0 0 0 0 0 0 0 0 0 0 0

Thaumastocoridae

Discocoris drakei 0 0 0 0 0 0 0 0 0 +++ 0 0 0

D. fernandezi 0 0 0 0 0 0 0 0 +++ 0 0 0 0

Pentatomidae

Lincus spp. ++ - 0 0 0 0 +++ +++ + 0 0 - -

0 absence, + less than 10 in an inflorescence, they are considered erratic, ++ between 10 and 100 in an inflorescence, +++ more than 100 in an inflorescence, - not confirmed.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

inflorescences, either during the day or at night.

However, Bernal and Ervik (1996) report sev-

eral species visiting the female inflorescences

of P. seemannii in Ecuador, but in much lower

numbers compared to the male inflorescences.

Syagrus spp.: This genus was not sampled

by us. We mention it here because it is the

exclusive host, along with the genus Oeno-

carpus, of the Derelomini Anchylorhynchus.

In Syagrus cocoides Mart., Syagrus vermicu-

laris Noblick, and Syagrus sancona (Kunth) H.

Karsten, Costa et al. (2009), de Melo-Valente

and Souza de Medeiros (2013), and Valente and

Madeiros, (2013) report many species of Cur-

culionidae Derelomini, including various spe-

cies of Anchylorhynchus (see below). Costa et al.

(2009), Silberbauer-Gottsberger et al. (2013), de

Melo-Valente and Souza de Medeiros (2013),

report different species in several species of

Syagrus (see below, insects). We found differ-

ent species of Anchylorhynchus in all the sampd

species of Oenocarpus. Therefore, this genus

of Derelomini is clearly dependent on Syagrus

and Oenocarpus.

The insects

Dynastidae (Coleoptera): Unlike other

common insect groups in neotropical palm

inflorescences that are highly specialized, such

as Curculionidae Derelomini, Nitidulidae Mys-

tropini, or Miridae Phylini, Dynastidae shows

no apparent specialization (Table 6). Adult

insects are attracted to palm inflorescences

from the moment the bract opens and anthesis

begins. Several authors have demonstrated that

they contribute to pollination, however, Mora-

Urpí and Solís (1980), Scariot et al. (1991), as

cited by Barfod et al. (2011), consider their role

to be secondary, with a limited function in pol-

len transfer. Moore and Jameson (2013) report

various unidentified Cyclocephala species on

several Arecaceae species. Aguiar-Gonçalves

(2018) studied the spatial distribution of Cyclo-

cephala in Brazil. Therefore, the role of Cyclo-

cephalini in pollination seems limited, and they

should be considered opportunists, similar to

Hymenoptera, Apidae (Table 6), which are very

common in palm inflorescences (Barfod et al.,

2011; Ervik et al., 1999).

• Ancognatha vulgaris in C. quindiuense

• C. amazona in B. gasipaes, O. bataua, and

M. flexuosa.

• P. macrocarpa ssp. P. tenuicaulis, E. oleifera.

• C. collaris in E.oleifera,

• C. couturieri on A. sp. aff. murumuru.

• C. discolor in A. urostachys, O. bataua, and

M. flexuosa

• C. distincta in E. oleifera, B. gasipaes, O.

mapora.

• C. goetzi in A. urostachys.

• C. guianae in M. flexuosa

• C. lunulata in A. sp. aff. murumuru.

• C. marginalis in A. urostachys, O. bataua

and Attalea microcarpa

• C. stictica in A. urostachys,

• C. variipenis in A. urostachys.

• M. signatoides in A. urostachys, O. bataua,

and O. mapora.

Some species are rare or observed in

small quantities: Caladenia marginata on A.

urostachys in Ecuador, associated with C. dis-

tincta and C. couturieri on A. aff. murumuru in

Uchiza, Peru (Dechambre, 1998).

Curculionidae (Coleoptera) (Table 2).

Many species are attracted to the anthesis of

most palm species. Most belong to the subfam-

ily Curculioninae, tribe Derelomini. The phy-

logeny of Derelomini was established by Franz

(2006). In a recent study, Haran et al. (2023) pro-

posed a synthesis on pollinating Curculionidae.

Not all genera and species mentioned in the

literature will be discussed here, only those with

confirmed generic and specific identification.

In Table 2, we report the most notable genera

observed by us, including references from other

authors. The genus Anchylorhynchus seems spe-

cialized in the genera Oenocarpus, Syagrus, and

Butia. We personally found it in all the sampled

species of Oenocarpus, including A. bicarinatus

O’Brien and Anchylorhynchus spp. (identifica-

tion in progress). Küchmeister et al. (1998)

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

Table 6

Presence of the Dynastidae: Cyclocephalinae: Ancognatha, Cyclocephala and Mimeoma in the inflorescences of palm trees.

Authors Collectors Genus, species Host plant

Country, province, department or state, locality

GC/CD/KM Ancognatha vulgaris Arrow Ceroxylon quindiuense Peru, San Martin, Ocol

Ervik et al. 1999 Cyclocephala aequatoria Endrödi Phytelephas seemannii Ecuador

KM/FK 1986

GC/FK 1992

FK/KM 10.1996

GC/FK 29.01.1994

C. amazona L. Astrocaryum macrocalyx Perú, Loreto, Jenaro Herrera,

Iquitos, Quistococha

FK 10.1985 C. amazona L Oenocarpus (Jessenia) bataua Perú, Loreto Iquitos

GC 13.10.1991 C. amazona L. Bactris gasipaes Perú, Loreto, Jenaro Herrera

GC/JV 13.4.91 C. amazona L. Mauritia flexuosa Perú, Loreto, Quistococha

GC/PB 18.10.2002

GC/FK 29.01.1992

C. amazona L. Elaeis oleifera

Astrocaryum macrocalyx

Ecuador, Quinindé

Perú, Iquitos

Bernal & Ervik,1996 Ervik et al. 1999, C. amazonica L. Phytelephas seemannii Ecuador

GC/JD 10.1993 C. collaris Burm. Elaeis oleífera Brasil, Bahia, Ilheus

GC/JLl10.1988 C. couturieri Dech. Astrocaryum aff. murumuru Perú, San Martin. Uchiza

GC/JLl 10.1988 C. dilatata Prell Astrocaryum macrocalyx Perú San Martin. Uchiza

LR/IS 02.09.1999 C. discolor Herbst Oenocarpus bataua Ecuador, Napo, Tena

LR/IS 02.11.1999 C. discolor Herbst Astrocaryum urostachys Ecuador, Napo, Tena

FK 10.04.1987 C. discolor Herbst Phytelephas macrocarpa Perú, San Martin, Uchiza

GC/JV 13.04.1991 C. discolor Herbst Mauritia flexuosa Perú, Loreto, Quistococha

Ervik et al. 1999 C. discolor Herbst Aphandra natalia Ecuador

Ponchel Y. http:// consult. 2015 C. discolor Herbst No identificado Perú

GC/FK 19.08.87 C. discolor Herbst No identificado Perú, San Martin Uchiza

LR 02.09.1999 C. discolor Herbst Jessenia bataua (=Oenocarpus) Ecuador, Napo, Tena

GC/JD 06.11.1993 C. distincta Burmeister Elaeis oleifera Brasil, Bahia Ilheus

GC/MSP 15.07.1997 C. distincta Burmeister Oenocarpus mapora Brasil, Belem Embrapa

GC/JD 06.02.1993 C. distincta Burmeister Bactris gasipaes Brasil, Bahia Ilheus

Bezerra de Souza et al. 2014 C. distincta Burmeister No identificado Brasil

GC/JD 11.1993 C. distincta Burmeister Attalea funifera Brasil, Bahia Ilheus

Moron et al. 2014 C. fasciolata Bates Astrocaryum mexicanum Mexico, Vera Cruz

Oliveira & Avila 2011 C. forsteri Endrödi Acronomia aculeata Brasil, Mato Grosso

T.P 19.11.2000 C. goetzi Endrödi Astrocaryum urostachys Ecuador, Remolino

GC/FK 05.1996 C. guyanae Dechambre Mauritia flexuosa Brasil, Amazonas, Manaus

GC/FK 05.1996 C. guyanae Dechambre Oenocarpus bacaba Brasil, Amazonas, Manaus

GC 09.1988 C. lunulate Burmeister Astrocaryum murumuru Perú, San Martin, Uchiza

TP 01.09.1999 C. marginalis Kirsch Oenocarpus bataua Ecuador, Tena

GC/FK 29.09.1991 C. marginalis Kirsch Phytelephas tenuicaulis Perú, San Martin, Uchiza

GC/FK 18.08.1987 C. peruana Endrödi Phytelephas tenuicaulis Perú, San Martin, Uchiza

Ervik et al. 1999 C. quadripunctata Hohne Phytephas macrocarpa Ecuador

TP 12.12.2000 C sticticaBurm. Astrocaryum urostachys Ecuador, Alto Pastaza

TP 19.09.2000 C. variipenis Miki Astrocaryum urostachys Ecuador, Pastaza Remolino

TP 05.11.2000 Mimeoma signatoides Höhne Oenocarpus bataua Ecuador, Pastaza Chunitayo

GC/FK 05.1996 Mimeoma maculata Burmeister. Oenocarpus bacaba Brasil, Amazonas, Manaus

TP 12.12.2000 Mimeoma signatoides Höhne Astrocaryum urostachys Ecuador, rio Pastaza

TP 09.09.2000 Mimeoma signatoides Höhne Oenocarpus bataua Ecuador, Chunitayo

CD/GC 06.10.2006 Mimeoma signatoides Höhne O. mapora Perú, rio Maniti, Villa Maria

GC/FK 05.1996 Mimeoma signatoides Höhne O. vacaba Brasil, Amazonas, Manaus

GC/PB/MSP 06.1997 Phileurus couturieri Dechambre O. mapora Brasil, Belem, Embrapa

GC 11.06.1992 Phileurus excavatus Prell Mauritia flexuosa Perú, Loreto, Jenaro Herrera, Fierrocaño

Collectors: PB: P. Beserra, GC: G. Couturier, JD: J. Delabie, CD: C. Delgado, FK: F. Kahn, KM: K. Mejía, MSP: M. Padilha,

TP: T. Peyret, LRL: L. Reynaud, and IS: I. Suarez.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

reported Anchylorhynchus tricarinatus Vaurie

on O. bataua. Souza de Medeiros and Núñez-

Avellaneda (2013) described three new species:

Anchylorhynchus pinocchio and Anchylorhyn-

chus luteobrunneus on S. sancona H. Karst.,

Anchylorhynchus centrosquamatus Medeiros &

Núñez on Syagrus orinocensis (Spruce) Burret

and reported that adults are pollinators while

the larvae develop in the endosperm, preventing

fruit development. Costa et al. (2009) reported

A. amazonicus Voss on S. cocoides Mart., and S.

vermicularis Noblick. Silberbauer- Gottsberger

et al. (2013) reported Anchylorhynchus camposi

Bondar on Syagrus petraea (Mart.) Becc., and

Anchylorhynchus bicolor Voss on Butia para-

guayensis (Barb.Rodr.) L.H.Bailey.

Guerrero-Oyala et al. (2018) reported sev-

eral unidentified species of Curculionidae on

S. sancona (Kunth) H. Karst., where the genus

Anchylorhynchus is undoubtedly found. Souza

de Medeiros and Núñez-Avellaneda (2013) cited

Anchylorhynchus elongatus on S. sancona, and

Souza de Medeiros and Vanin (2020) described

Anchylorhynchus multisquamis on Syagrus sp.

Souza de Medeiros et al. (2019) reported Anchy-

lorhynchus trapezicollis on Syagrus coronata,

and de Melo-Valente and Souza de Medeiros

(2013) described Anchylorhynchus vanini on

S. vermicularis Noblick. Guerrero-Oyala and

Núñez-Avellaneda (2017) reported A. luteob-

runneus on S. sancona (Kunth) H. Karst., Souza

de Medeiros and Nuñez-Avellaneda (2013)

reported A. elongatus also on S. sancona, and

Medeiros and Farrell (2019) described A. tra-

pezicollis on S. coronata. Souza de Medeiros

and Vanin (2020) described A. multisquamis on

Syagrus sp. Souza de Medeiros et al. (2019) and

Haran et al. (2023) reported that females lay

their eggs in the female flowers, thereby limit-

ing their role in fruit production (Silberbauer-

Gottsberger et al., 2013).

The genus Andranthobius: de Melo-Valente

and Lima da Silva (2014) reported Andran-

thobius setirostris on S. cocoides, and Mora-

Urpí and Mexzon (1996) on A. palmarum

in B. gasipaes.

The genus Phyllotrox is cited by most

authors, including ourselves (six different

species separated in our collections, (O’Brien,

pers. comm.), on most palm genera and spe-

cies. Listabarth (1996) considers them to be

pollinators (Table 2) The specific identifica-

tion of this genus is extremely difficult, but

Küchmeister et al. (1998) cited ten species, and

Voisin (1989) described Phyllotrox lamottei in

Venezuela without specifying the host plant.

We found an undetermined species of

Derelomini with red larvae, which was very

abundant in all the sampled P. tenuicaulis and

entirely absent from other palms.

The genus Celetes includes several species

considered to be pollinators; however, they

should also be regarded as parasites of young

fruits (Table 1). An unidentified species was

found exclusively on A. funifera in the Atlantic

Forest (Ilheus, Brazil).

The genus Terires, with Terires minusculus,

was found only on Astrocaryum of the subgenus

Pleiogynanthus. It develops in young fruits, and

in a count of 200 fruits, a parasitization rate of

35 % was revealed.

Nitidulidae (Coleoptera): The tribe Mys-

tropini (Table 2) was revised by Kirejtshuk

& Couturier (2010), where ten new species

were described. They are considered the most

effective pollinators, alongside Curculionidae

Derelomini. Núñez-Avellaneda et al. (2005)

studied the role of climate in pollination by

Mystropini. Scariot et al. (1991) reported Mys-

trops cf. minusculus on A. aculeata (not studied

by us). Restrepo-Correa et al. (2016) reported

a very high degree of specialization between

Mystrops and Wettinia spp. in Colombia.

Since the Mystrops were not identified at the

species level, the relationship with other palm

genera and species cannot be established. How-

ever, these results confirm a high level of spe-

cialization, as seen in the review by Kirejtshuk

& Couturier (2010).

Staphylinidae (Coleoptera): The genus

Arecaceopora Pace appears to depend on P.

tenuicaulis, with three species: A. couturieri,

Arecaceopora delgadoi, and Arecaceopora per-

lifera (Pace, 2010) (Table 2). Interestingly, in

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

Ecuador, it is the genus Amazoncharis that is

found on P. seemanni (Bernal & Ervik, 1996;

Ervik et al., 1999). Oussipaliaglossa couturieri

Pace was found only on A. funifera (outside the

Amazon), and D. amazonensis, a unique spe-

cies on M. flexuosa, was very abundant on all

male inflorescences of the sampled individuals

(Chani-Posse & Couturier, 2012).

Heteroptera (Table 5) are characterized

by three main families. The Miridae Phylini,

with the genera Alvarengamiris (A. alvarengai

Carvalho, A. kemberi Costa & Couturier, A.

margaridae Costa & Couturier, are found only

in the male inflorescences of M. flexuosa and

M. carana, where they are highly abundant and

reproduce. The Anomalocornis genus includes

six known species in palms: A. couturieri Carv-

alho & Costa, Anomalocornis geijskesi Carvalho

& Wygodzinsky, Anomalocornis gentyi Costa

& Couturier, Anomalocornis peyreti Couturier

& Costa, Anomalocornis rondoniensis Carval-

ho, and Anomalocornis tucuruiensis Carvalho,

which are associated with Astrocaryum spp. and

B. gasipaes (Carvalho & Costa, 1999; Couturier

et al., 2003; Costa & Couturier, 2002; Costa

& Couturier, 2012). The host plant for Astro-

caryum ariasi Carvalho from Manaus (Brazil)

is unknown, though it is likely a palm that

remains to be identified. Parafulvius henryi

Costa & Couturier is found on A. urostachys

(Costa & Couturier, 2000), and Fulvius chaque-

nus Carvalho & Costa on A. javarense. Lis-

tabarth (1996) reported the presence of an

unidentified Miridae on B. gasipaes.

The Thaumastocoridae (Heteroptera)

family includes two highly specialized spe-

cies of Discocoris: D. drakei Slater & Ashlock

on Oenocarpus spp. (Couturier et al., 2002;

Núñez-Avellaneda et al., 2015) and Discocoris

fernandezi Slater & Brailowsky on M. flexuosa

and M. carana (Couturier et al., 1998; Coutu-

rier, et al., 2002).

The Pentatomidae (Heteroptera) fam-

ily includes the genus Lincus, which has been

found on Astrocaryum spp. and E. guineensis,

where they are known to transmit wilt dis-

ease (Couturier & Kahn, 1989a; Couturier &

Kahn, 1989b; Llosa et al., 1990; Rolston, 1983;

Rolston, 1989), as well as Antiteuchus kerzhneri

Rider (2006) on M. flexuosa.

Apidae (Hymenoptera): Several species

of Apidae visit palm inflorescences without

any particular specialization (Table 4). These

are primarily Trigoninae and Apis mellifera.

Campbell et al. (2018) and Bezerra et al. (2020)

observed that Apidae are the most important

pollinators for E. oleracea.

On P. seemanni in Ecuador: Melipona fasci-

ata Cockerell, Trigona amalthea Olivier, Trigona

chanchamayoensis Schwarz, Trigona ferricauda

Cockerell, Trigona hyalinata Lepeletier, Trigona

nigerrima Cresson, Trigona schultesii Friese,

Trigona spinipes Fab, Trigona williana Friese,

and Trigona sp. (Bernal & Ervik, 1996).

Küchmeister et al. (1998) report Ptilotri-

gona lurida Smith on Astrocaryum acaule, Tri-

gona branneri Cockerell cf. on Astrocaryum

gynacanthum and A. attaleoides, as well as T.

williana Friese and Trigona fulviventris guianae

on A. microcarpa.

Other insects: There are many other

insects. Núñez-Avellaneda (2014) found 79

species as occasional or permanent visitors to

O. bataua, O. balickii, and Oenocarpus minor,

contributing to pollination or participating in

the degradation (e.g., Ptiliidae in P. te n ui c au -

lis, Couturier G and Delgado C, pers. obs.) of

the inflorescences. For example, unidentified

larvae of Drosophilidae develop in the male

inflorescences of P. tenuicaulis. The Droso-

philidae Palmophila ecuadoriensis and Palmo-

myia incerta were described by Grimaldi et al.

(2003) on Wettinia maynensis (P.e.), Chamae-

dorea linearis, and P. seemannii (P.i.). Many

species of Hymenoptera (Formicidae, Vespi-

dae), Diptera, and Chrysomelidae beetles have

been reported in various palm inflorescences

(Küchmeister et al., 1998).

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

DISCUSSION

The detailed analysis of the fauna associ-

ated with various genera and species of neo-

tropical palms, particularly regarding the fauna

of their inflorescences, reveals different levels

of specialization or coevolution. It goes without

saying that only precise species identification

allows the linkage between insect species and

the relevant genera and species of palms. This

has not always been possible.

Samples were taken outside the Amazo-

nian region, in the Peruvian Andes at 2 300 m

of altitude on C. quindiuense, as well as in the

Brazilian Atlantic Forest on A. funifera. This

demonstrates that geographic factors are not

the primary drivers of species distribution;

instead, the genera and species of palms deter-

mine the presence of different insect species.

One family (tribe) of insects and

one genus or subfamily of palms (as in the

case of Astrocaryum ssg., Monogynanthus

and Pleiogynanthus).

One insect species is linked to one or two

palm species (within the limits of our current

knowledge). Therefore, we can consider this

as coevolution. This specialization is strictly

related to the specific genera and species of

palms in question. In fact, the collections made

outside the Amazon, both from Ceroxylon in

the Peruvian Andes and from A. funifera in the

Brazilian Atlantic Forest, have demonstrated

that geography does not play a major role in

the distribution of insect species. Godsoe et

al. (2009) showed that the mutualism between

insect and plant does not depend on climatic or

geographical factors, but rather on the presence

of the host plant species.

Some examples: The Curculionidae Dere-

lomini Couturierius and Grasidius live exclu-

sively on E. oleifera (O’Brien et al., 2004). They

are replaced by several species of Elaeidobius,

imported from Africa, on E. guineensis, which

was also imported from Africa. M. flexuosa

and M. carana host a highly specialized fauna,

such as the Miridae of the genus Alvarengamiris

(three species), the Mystropini M. dalmasi,

the Thaumastocoridae D. fernandezi, and the

Staphylinidae D. amazonensis. The Nitidulidae

Mystropini Mystrops nitidulus is present in the

genus Attalea (A. funifera in the Atlantic For-

est of Brazil and A. plowmanii in the Iquitos

region, Allpahuayo Reserve), as reported by

Núñez-Avellaneda et al. (2005). However, it is

also found in Syagrus but is completely absent

from other studied palm genera.

The highest level of specialization is

observed at the tribal level (Derelomini for Cur-

culionidae, Mystropini for Nitidulidae, Philini

for Miridae). It is highly likely that the chemi-

cal compounds emitted by palms, as shown

by Knudsen (1999) for the palms Geonomeae

and Lajis et al. (1985) for E. guineensis, as well

as thermogenesis (Moore & Jameson, 2013),

determine the selective attraction of palms

to insects. Additionally, Godsoe et al. (2009)

showed that the mutualism between insect and

plant does not depend on climatic factors, but

rather on the presence of the host plant.

The increase in the temperature of spathes

before their opening has been noted by several

authors: Schroeder (1978), Küchmeister et al.

(1998), and more recently Pincebourde et al.

(2016). This phenomenon contributes to the

attraction of insects, as observed in O. mapora.

In this study, the role of insects in pollina-

tion is discussed. It is important to note that

pollination is only confirmed if the insects in

question actually move from the male to the

female flowers and carry viable and compat-

ible pollen (Beserra, 2002). For example, in the

case of M. flexuosa, Khorsand-Rosa and Koptur

(2013) demonstrated that insects play no role

in its pollination, a fact we also observed: the

abundant fauna found on the male inflores-

cences is absent from the female inflorescences.

However, this observation is questioned by

Mendes et al. (2016). In the female inflores-

cences, we only observed insects that were

“predators” of the fruits, such as Tyrannion sp.

(Curculionidae), several species of Baridinae,

and Leptoglossus hesperus (Coreidae) (Vasquez

et al., 2008). It should also be emphasized that

Baridinae are generally rare or absent during

anthesis and only approach the inflorescences

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026183, enero-diciembre 2026 (Publicado May. 15, 2026)

when the young fruits are forming, as seen with

Palmelampius heinrichi on B. gasipaes (O’Brien

& Kovarik, 2000).

Some groups of insects considered to be

pollinators are ubiquitous, such as Cyclocephala

spp. (Dynastidae) and bees of the genus Tri-

gona sensu lato, which visit various palm spe-

cies indiscriminately during anthesis. However,

Campbell (2018) consider native bees to be the

main pollinators of E. oleracea in Brazil.

Thus, many studies leave uncertain-

ty regarding pollination (Meekijjaroenroj &

Anstett, 2003). Barfod et al. (2011) and Nuñez-

Avellaneda et al. (2005) point out that experi-

ments to control pollination have rarely been

carried out, and that anemophily likely plays

a more important role than previously antici-

pated (Rios et al., 2014). In fact, it should be

considered that the insect in question must

carry viable pollen and visit the female flowers.

This point is highlighted by Núñez-Avellaneda

et al. (2015), and it is also important that the

insects do not develop on the young fruits

(Beserra, 2002).

Regarding the close relationship between

insects and their host plants, it will be necessary

to better understand the volatile compounds

of palm inflorescences, as demonstrated by

Lajis et al. (1985) for E. guineensis, as well as

advancements in the specific identification of

insects. We encountered difficulty in identify-

ing certain groups, such as Nitidulidae Mystro-

pini and Curculionidae Derelomini. In fact, too

often some authors label species as sp. A, B, etc.,

or sp. 1, 2, which only introduces confusion

between different studies. All these examples

confirm the close relationship between cer-

tain insect groups and their host palms, and

these conclusions are consistent with those

of Haran et al. (2023) for Curculionidae and

Kergoat et al. (2006) which show the phyloge-

netic relationships between the genus Acacia

(Fabaceae: Mimosoideae) and its associated

seed predators.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

To the Institute of Amazonian Research

of Peru (IIAP) and the Institute of Research

for Development (IRD, formerly ORSTOM)

for funding this research. To Marcos Garcia

(Embrapa, Manaus, Brazil), Giovanni Hon-

oré, director of the Otonga Reserve (Ecuador),

Maria Socorro Padilha de Oliveira (Embra-

pa, Belem), Wilson Gonzalez (IIAP, Iquitos),

Thomas Peyret and Lauriane Reynaud (Paris),

Ivan Suarez (Ecuador). Also, to the taxonomists

who identified and described the insects: Harry

Brailowsky (University of Mexico), Luis Costa

(National Museum of Rio de Janeiro, Bra-

zil), Roger Paul Dechambre (Museum, Paris),

Charles O’Brien (Arizona, United States of

America), Roberto Pace (Verona, Italy), Hélène

Perrin (Museum, Paris), Alexander Kirejtshuk

(University of St. Petersburg, Russia), Claus

Rasmussen (University of Aarhus, Denmark),

L.H. Rolston (University of Baton Rouge, Loui-

siana, United States of America), B. de Medeiros

(Field Museum of Natural History, Chicago,

United States of America). A special thanks to

Francis Kahn, botanist of the IRD, specialist

in neotropical palms, who has always followed

this work with interest and contributed to the

collections. Many thanks also to Paulo Beserra

(Embrapa, Belem) that participated with the

authors in many collects in Brazil and Ecua-

dor, and to all the farmers and villagers from

the various surveyed locations for their help

in the field.

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