Dahmer, Priscila C. ¹ ; Cavalcante, Ana Cristina C. ² and Demite, Peterson R. ³

¹Instituto de Ciências Exatas e Tecnologia, Universidade Federal do Amazonas, UFAM, Itacoatiara, Amazonas, Brazil.
²Instituto de Ciências Exatas e Tecnologia, Universidade Federal do Amazonas, UFAM, Itacoatiara, Amazonas, Brazil.
³✉ Campus Bonfim, Instituto Federal de Roraima, IFRR, Bonfim, Roraima, Brazil.

2026 - Volume: 66 Issue: 1 pages: 35-49

Original research

Keywords

Abstract

Introduction

Understanding mite fauna associated with agricultural environments is fundamental for developing effective management strategies for these ecosystems (Feres et al. 2005). Although most studies on mite diversity in agroecosystems have been carried out in southeastern Brazil (Pallini et al. 2007), much remains unknown, particularly regarding the acarofauna linked to crop systems in other regions. Studies on mite diversity are essential because, while several species are agricultural pests, many others contribute significantly to biological control—either by preying on phytophagous mites, insects, and weeds, or by serving as alternative prey for various predators (Moraes et al. 2024). Among these beneficial groups, mites of the family Phytoseiidae are considered as key predators of pest mites and small insects in agroecosystems worldwide (McMurtry et al. 2013)

In the state of Amazonas, research on mite fauna in agricultural systems is still scarce and relatively recent, with emphasis on citrus and palm crops. Bobot et al. (2011) reported 25 mite species in an orange orchard in Manaus, including ten belonging to the Phytoseiidae family. Nuvoloni et al. (2015a) identified 11 Phytoseiidae species in rubber plantations (Hevea brasiliensis Muell. Arg., Euphorbiaceae). Ferreira et al. (2018) recorded 32 mite species—eight of which were Phytoseiidae—in citrus crops in Iranduba and Manaus, in addition to 25 species (nine Phytoseiidae) found on spontaneous vegetation. Cruz et al. (2015) investigated mites associated with Raoiella indica Hirst (Tenuipalpidae) in coconut palms across four municipalities and reported 18 phytoseiid species. Later, Cruz et al. (2019) analyzed oil palm systems (Elaeis guineensis Jacq., E. oleifera (Kunth) Cortés, and hybrids), reporting 6, 17, and 3 phytoseiid species, respectively. The most comprehensive study was conducted by Vasconcelos and Silva (2015), who recorded mites on 101 plant species, identifying Phytoseiidae as the dominant predators.

Phytoseiidae includes several species of economic importance as biological control agents (McMurtry et al. 2013; Knapp et al. 2018). In Brazil, approximately 260 species have been recorded, with 55 of these reported in the state of Amazonas (Lofego et al. 2024; Demite et al. 2025). The present study aims to assess the diversity of Phytoseiidae associated with guarana (Paullinia cupana Kunth, Sapindaceae), a native Amazonian crop, in central Amazonia.

Material and methods

Sampling areas

The study was conducted in 20 areas across six municipalities in the central part of the Brazilian Amazonian biome, most of which were managed under agroecological or low-input systems and surrounded by continuous native forest: Itacoatiara (five areas), Urucará (four), Itapiranga (two), Maués (four), São Sebastião do Uatumã (three), and Urucurituba (two) (Figure 1; Table 1).

The climate of the study region is classified as'Af' according to the Köppen-Geiger system, with two distinct seasons: a rainy season beginning in December and a less rainy season starting in May. Rainfall is significant year-round, with an annual total of 2,261 mm. September is the driest month and March the wettest. The average annual temperature is 26.9 °C (Climate-Data 2025).

Sampling

Sampling was conducted at irregular intervals from May 2019 to January 2021, encompassing both the rainy and less rainy seasons, according to site accessibility and logistical constraints. Each guarana plantation area was sampled one to four times during the study period. All guarana plantations surveyed were located within forested landscapes and surrounded by native forest and/or secondary vegetation under natural regeneration, ensuring strong connectivity with adjacent natural habitats.

In each sampling event, 10 leaves were collected from each of 15 randomly selected plants per site, totaling 150 leaves per event. Across the 27 sampling events conducted in this study, a total of 4,050 leaves were sampled and processed for mite extraction. Leaves were collected using pruning shears and were placed in paper bags, sealed inside polyethylene bags, and stored in insulated polystyrene boxes with ice.

Collected material was examined under a stereomicroscope (Leica EZ4, 40x magnification). The phytoseiid mites found were carefully removed from the leaves using a fine-bristled brush and mounted on microscope slides in Hoyer's medium (Krantz and Walter, 2009). Slides were kept in an oven at 50–60 °C for approximately three days. Afterwards, the edges of the coverslips were sealed with clear nail polish.

Mite identification

Specimens were identified under a phase-contrast optical microscope (Zeiss Axio Imager.M3). Identification at the genus level followed Chant and McMurtry (2007), while species-level identification was based on the available literature (e.g., Lofego et al. 2024, and references cited for each identified species). Only adult specimens of each species were considered, since most species cannot be reliably identified from immature stages.

Data analysis

To summarize the mite community structure, species-site interactions were visualized using a chord diagram generated with the RAWGraphs open-source platform (Mauri et al. 2017). This visualization represents the distribution and connectivity of the Phytoseiidae community across the 20 sampled localities, where the thickness of the arcs is proportional to the total abundance of each species-site association.

Voucher specimen deposition

Voucher specimens were deposited in the mite collection of the Acarology Laboratory, Department of Biological Sciences, São Paulo State University (UNESP), São José do Rio Preto, São Paulo State, Brazil.

Results

A total of 408 specimens of phytoseiid mites were collected in this study (Table 2). The recorded species belong to 11 genera, across two subfamilies: Amblyseiinae, with 19 species, and Typhlodrominae, with four (Table 2). The genera Amblyseius and Iphiseiodes exhibited the highest species richness, with eight and four species, respectively. Among the 23 phytoseiid species recorded, four are probably new—three from the genus Amblyseius and one from Proprioseiopsis. The most abundant species was Typhlodromips angustus Guanilo & Moraes, with 172 specimens collected (42.2% of the total abundance), followed by Proprioseiopsis neotropicus (Ehara) (48 specimens), Iphiseiodes zuluagai Denmark & Muma (39), and Amblydromalus itacoatiarensis Demite, Cavalcante & Lofego (32) (Figure 3). The species found in the greatest number of sites were P. neotropicus, T. angustus, A. itacoatiarensis, and Amblyseius aerialis (Muma), present in 13, 12, 8, and 8 sites, respectively.

The interaction network between phytoseiid species and sampling sites in guarana cultivation revealed a structured community, composed of both broadly distributed species and taxa with restricted occurrence (Figure 3). Typhlodromips angustus and P. neotropicus were the most connected and abundant species, suggesting high ecological plasticity and a central role in the natural regulation of mite pests in the Amazon region. Conversely, less frequent species were limited to one or few locations, indicating dependence on specific microhabitat conditions in guarana agroecosystems.

Amblyseiinae

Amblydromalus itacoatiarensis Demite, Cavalcante & Lofego

Amblydromalus itacoatiarensis Demite, Cavalcante and Lofego, 2019: 2484.

Specimens examined — Itacoatiara (Area 3): 02-VII-19 (2); 10-X-19 (1); 06-I-20 (4); Itacoatiara (Area 4): 08-VIII-19 (11); Itapiranga (Area 1): 04-XII-19 (1); Itapiranga (Area 2): 04-XII-19 (3); Maués (Area 4): 11-III-20 (1); São Sebastião do Uatumã (Area 3): 17-XI-20 (1); Urucará (Area 2): 06-XI-19 (3); Urucurituba (Area 1): 21-XII-20 (5).

Amblyseius aerialis (Muma)

Amblyseiopsis aerialis Muma, 1955: 264.

Amblyseius aerialis – Athias-Henriot, 1957: 338.

Typhlodromus (Amblyseius) aerialis – Chant, 1959: 88.

Amblyseius (Amblyseius) aerialis – Denmark and Muma, 1989: 15.

Specimens examined — Itacoatiara (Area 1): 14-X-19 (1); Itapiranga (Area 1): 04-XII-19 (1); Itapiranga (Area 2): 04-XII-19 (1); São Sebastião do Uatumã (Area 1): 17-XI-20 (2); São Sebastião do Uatumã (Area 2): 17-XI-20 (1); São Sebastião do Uatumã (Area 3): 17-XI-20 (1); Urucará (Area 3): 06-XI-19 (2); Urucará (Area 4): 06-XI-19 (4).

Amblyseius chiapensis De Leon

Amblyseius chiapensis De Leon, 1961: 85.

Amblyseius triplaris De Leon, 1967: 25 (synonymy according to Denmark and Muma, 1989).

Specimens examined — Urucará (Area 4): 06-XI-19 (1).

Amblyseius duckei Nuvoloni, Lofego, Rezende & Feres

Amblyseius duckei Nuvoloni, Lofego, Rezende and Feres, 2015a: 191.

Specimens examined — Itapiranga (Area 1): 04-XII-19 (2).

Amblyseius manauara Nuvoloni, Lofego, Rezende & Feres

Amblyseius manauara Nuvoloni, Lofego, Rezende and Feres, 2015a: 191.

Specimens examined — Itacoatiara (Area 2): 25-VI-19 (3); 07-X-19 (4); Itacoatiara (Area 3): 02-VII-19 (1); Itacoatiara (Area 5): 17-X-19 (2); Urucará (Area 3): 06-XI-19 (2).

Amblyseius vasiformis Moraes & Mesa

Amblyseius vasiformis Moraes & Mesa, in Moraes et al., 1991: 119.

Specimens examined — Itacoatiara (Area 3): 06-I-20 (2); Urucará (Area 3): 06-XI-19 (3).

Amblyseius sp. 1

Specimens examined — Itacoatiara (Area 1): 08-I-20 (1); Maués (Area 2): 11-III-20 (1).

Amblyseius sp. 2

Specimens examined — Itacoatiara (Area 2): 07-X-19 (2); 10-I-20 (3); Itacoatiara (Area 3): 10-X-19 (1); Itacoatiara (Area 5): 17-X-19 (3); São Sebastião do Uatumã (Area 1): 17-XI-20 (1); Urucará (Area 2): 06-XI-19 (1).

Amblyseius sp. 3

Specimens examined — Itacoatiara (Area 2): 25-IV-19 (2); Itacoatiara (Area 3): 02-VII-19 (2).

Arrenoseius urquharti (Yoshida-Shaul & Chant)

Amblyseius urquharti Yoshida-Shaul and Chant, 1988: 2055.

Fundiseius urquharti – Moraes et al., 2004: 89.

Specimens examined — Itacoatiara (Area 1): 08-I-20 (3).

Iphiseiodes kamahorae De Leon

Iphiseiodes kamahorae De Leon, 1966: 84.

Specimens examined — Itacoatiara (Area 1): 22-V-19 (3); Itacoatiara (Area 3): 02-VII-19 (1); 06-I-20 (1); Itacoatiara (Area 4): 08-VII-19 (1); Itapiranga (Area 1): 04-XII-19 (6); Maués (Area 4): 11-III-20 (2); Urucará (Area 1): 06-XI-19 (1).

Iphiseiodes katukina Nuvoloni, Lofego, Rezende & Feres

Iphiseiodes katukina Nuvoloni, Lofego, Rezende and Feres, 2015a: 195.

Specimens examined — Itacoatiara (Area 1): 22-V-19 (3); 14-X-19 (1); 08-I-20 (2); Maués (Area 2): 11-III-20 (2); Urucará (Area 3): 06-XI-19 (3); Urucurituba (Area 2): 21-XII-20 (3).

Iphiseiodes raucuara Nuvoloni, Lofego, Rezende & Feres

Iphiseiodes raucuara Nuvoloni, Lofego, Rezende and Feres, 2015a: 195.

Specimens examined — Itacoatiara (Area 1): 08-I-20 (1); Itacoatiara (Area 2): 06-I-20 (1); 25-VI-19 (1); Itapiranga (Area 2): 04-XII-19 (1); São Sebastião do Uatumã (Area 1): 17-XI-20 (1).

Iphiseiodes zuluagai Denmark & Muma

Iphiseiodes zuluagai Denmark and Muma, 1972: 23.

Amblyseius zuluagai — Moraes and Mesa, 1988: 79.

Specimens examined — Itacoatiara (Area 2): 25-VI-19 (17); 07-X-19 (7); 10-I-20 (6); Itacoatiara (Area 5): 17-X-19 (1); São Sebastião do Uatumã (Area 1): 17-XI-20 (6); Urucará (Area 2): 06-XI-19 (1); Urucará (Area 4): 06-XI-19 (1).

Paraamblyseius multicircularis Gondim Jr. & Moraes

Paraamblyseius multicircularis Gondim Jr. and Moraes, 2001: 79.

Specimens examined — Maués (Area 3): 11-III-20 (1).

Proprioseiopsis neotropicus (Ehara)

Amblyseius neotropicus Ehara, 1966: 133.

Proprioseiopsis neotropicus — Moraes et al., 1986: 119.

Specimens examined — Itacoatiara (Area 1): 22-V-19 (8); Itacoatiara (Area 2): 25-IV-19 (1); Itacoatiara (Area 3): 02-VII-19 (6); Itacoatiara (Area 4): 08-VIII-19 (3); Itapiranga (Area 1): 04-XII-19 (3); Itapiranga (Area 2): 04-XII-19 (2); Maués (Area 1): 11-III-20 (1); Maués (Area 4): 11-III-20 (2); São Sebastião do Uatumã (Area 3): 17-XI-20 (6); Urucará (Area 1): 06-XI-19 (7); Urucará (Area 2): 06-XI-19 (1); Urucará (Area 3): 06-XI-19 (7); Urucurituba (Area 1): 21-XII-20 (1).

Proprioseiopsis ovatus (Garman)

Amblyseiopsis ovatus Garman, 1958: 78.

Typhlodromus (Amblyseius) ovatus — Chant, 1959: 90.

Proprioseiopsis ovatus — Denmark and Muma, 1973: 237.

Proprioseiopsis (Proprioseiopsis) ovatus — Karg, 1989: 208.

Amblyseiulus cannaensis Muma, 1962: 4 (synonymy according to Denmark and Evans, 2011).

Proprioseiopsis cannaensis — Muma et al., 1970: 38.

Proprioseiopsis (Proprioseiopsis) cannaensis — Karg, 1989: 116.

Amblyseiulus hudsonianus Chant and Hansell, 1971: 723 (synonymy according to Denmark and Evans, 2011).

Amblyseius parapeltatus Wu and Chou, 1981: 274 (synonymy according to Tseng, 1983).

Amblyseius peltatus Van der Merwe, 1968: 119 (synonymy according to Tseng, 1983).

Amblyseius (Proprioseiopsis) peltatus — Blommers, 1976: 100.

Proprioseiopsis peltatus — Moraes et al., 1986: 121.

Iphiseius punicae Gupta, 1980: 213 (synonymy according to Gupta, 1985).

Proprioseiopsis punicae — Moraes et al., 1986: 122.

Specimens examined — São Sebastião do Uatumã (Area 1): 17-XI-20 (1); Urucará (Area 3): 06-XI-19 (2).

Proprioseiopsis sp.

Specimens examined — Itacoatiara (Area 2): 25-VI-19 (2); 10-I-20 (1); Itapiranga (Area 1): 04-XII-19 (4); Itapiranga (Area 2): 04-XII-19 (2); Urucará (Area 4): 06-XI-19 (11).

Typhlodromips angustus Guanilo & Moraes

Typhlodromips angustus Guanilo & Moraes, in Guanilo et al., 2008: 34.

Specimens examined — Itacoatiara (Area 1): 22-V-19 (3); 14-X-19 (22); 08-I-20 (25); Itacoatiara (Area 3): 02-VII-19 (1); 10-X-19 (12); 06-I-20 (15); Itacoatiara (Area 4): 08-VIII-19 (1); Itapiranga (Area 2): 04-XII-19 (2); Maués (Area 1): 11-III-20 (2); Maués (Area 2): 11-III-20 (16); Maués (Area 4): 11-III-20 (4); São Sebastião do Uatumã (Area 2): 17-XI-20 (21); Urucará (Area 1): 06-XI-19 (11); Urucará (Area 2): 06-XI-19 (23); Urucurituba (Area 1): 21-XII-20 (6); Urucurituba (Area 2): 21-XII-20 (8).

Typhlodrominae

Cocoseius palmarum Gondim Jr., Moraes & McMurtry

Cocoseius palmarum Gondim Jr., Moraes and McMurtry, 2000: 1226.

Specimens examined — Itacoatiara (Area 2): 25-VI-19 (1).

Galendromimus (Galendromimus) alveolaris (De Leon)

Typhlodromus alveolaris De Leon, 1957: 141.

Typhlodromus (Typhlodromus) alveolaris — Chant, 1959: 52.

Galendromimus alveolaris — Muma, 1961: 297.

Cydnodromella alveolaris — Chant and Yoshida-Shaul, 1986: 2820.

Galendromimus (Galendromimus) alveolaris — Chant and McMurtry, 1994: 242.

Specimens examined — Itacoatiara (Area 3): 10-X-19 (1).

Leonseius regularis (De Leon)

Typhloseiopsis regularis De Leon, 1965: 122.

Diadromus regularis — De Leon, 1966: 100.

Chanteius regularis — De Leon, 1967: 16.

Typhlodromus regularis — Chant and Yoshida-Shaul, 1983: 1034.

Leonseius regularis — Chant and McMurtry, 1994: 258.

Specimens examined — Urucará (Area 3): 06-XI-19 (2).

Metaseiulus (Metaseiulus) adjacentis (De Leon)

Typhlodromus adjacentis De Leon, 1959: 124.

Typhlodromina adjacentis — Muma, 1961: 297.

Paraseiulella adjacentis — Denmark, 1994: 18.

Metaseiulus (Metaseiulus) adjacentis — Moraes et al., 2000: 256.

Specimens examined — Itacoatiara (Area 2): 10-I-20 (1); São Sebastião do Uatumã (Area 1): 17-XI-20 (1).

Discussion

Our study provides the first comprehensive assessment of the phytoseiid mite fauna associated with guarana cultivation in northern Brazil, revealing a high diversity of species. This assemblage not only highlights the largely unexplored biodiversity of the Brazilian Amazon but also reinforces the importance of agricultural ecosystems as reservoirs of predatory species that contribute to essential ecosystem services, particularly the natural regulation of herbivorous pests. Importantly, most of the sampled plantations were managed under agroecological or low-input systems and exhibited high internal environmental heterogeneity, with diverse understory vegetation and mixed plant strata. In addition, all sites were surrounded by continuous or semi-continuous native forest, ensuring strong connectivity with natural habitats — a landscape configuration rarely observed in crop systems of southern and southeastern Brazil, which are typically embedded in highly fragmented and intensively managed agricultural matrices. More structurally complex or diversified agroecosystems, which incorporate intercropping, ground cover, or are adjacent to semi-natural or natural habitats, can function as reservoirs for predatory mites, enhancing species richness and enabling their persistence and dispersal across agricultural landscapes (Tixier 2018).

Remarkably, five species reported in this survey have only recently been described (last ten years), further emphasizing the lack of historical data and the urgency of continued taxonomic exploration in the region: Amblydromalus itacoatiarensis Demite, Cavalcante & Lofego; Amblyseius duckei Nuvoloni, Lofego, Rezende & Feres; Amblyseius manauara Nuvoloni, Lofego, Rezende & Feres; Iphiseiodes katukina Nuvoloni, Lofego, Rezende & Feres; and Iphiseiodes raucuara Nuvoloni, Lofego, Rezende & Feres (Nuvoloni et al. 2015a; Demite et al. 2019). Similar patterns of records of recently described species or potential new taxa have been reported in other crops in Amazonian Region, including citrus (Bobot et al. 2011; Ferreira et al. 2018), açaí palm (Brito et al. 2024) and cocoa (Brito et al. 2025). The presence of recently described taxa suggests that the diversity of predatory mites in Amazonian agroecosystems is still minimally documented.

Such a concentration of recently recognized taxa within a single cropping system is unusual in agricultural environments of other Brazilian regions, where mite communities are typically dominated by well-known generalist species. Previous surveys in perennial crops in other regions of Brazil, mainly in the South and Southeast (e.g., Hernandes and Feres 2006; Daud and Feres 2013; Reichert et al. 2014; Mineiro and Raga 2020; Araújo et al. 2022; Ferla et al. 2023; Melo et al. 2023) have rarely reported comparable numbers of newly described phytoseiids or potential undescribed species. This pattern suggests that Amazonian agroecosystems continue to reveal undocumented components of regional phytoseiid fauna.

Furthermore, our survey expands the known distribution of two species reported for the first time in the state of Amazonas: Iphiseiodes raucuara, previously reported from Acre and Mato Grosso (Nuvoloni et al. 2015a; Demite et al. 2021), and Galendromimus (Galendromimus) alveolaris, previously recorded in the states of Bahia, Mato Grosso, Mato Grosso do Sul, and São Paulo (Moraes et al. 1993; Feres and Moraes 1998; Zacarias and Moraes 2001; Ferla and Moraes 2002; Daud and Feres 2005; Demite et al. 2011, 2012, 2013; Mendonça et al. 2019). These findings reinforce that the Amazonian phytoseiid fauna remains underrepresented in distributional records.

Several species detected in this study, including Amblyseius aerialis, Amblyseius chiapensis, Iphiseiodes zuluagai, Proprioseiopsis neotropicus, and Proprioseiopsis ovatus, are widely distributed across Brazil (Feres and Moraes 1998; Ferla and Moraes 2002; Gondim Jr and Moraes 2001; Zacarias and Moraes 2001; Feres et al. 2005; Buosi et al. 2006; Demite et al. 2009, 2011, 2017, 2021; Rezende et al. 2011; Gonçalves et al. 2015; Nuvoloni et al. 2015a,b; Rocha et al. 2015; Lofego et al. 2017, 2024; Mendonça et al. 2019; Cavalcante et al. 2021). The high abundance of P. neotropicus and I. zuluagai as the second and third most dominant species indicates their ecological resilience. Their coexistence alongside regionally adapted taxa such as T. angustus (the most abundant) and A. itacoatiarensis (the fourth most abundant) suggests a functionally diverse predator assemblage. A similar dominance of A. aerialis and I. zuluagai as key predatory mites has also been documented in other perennial crops within the Central Amazon. Ferreira et al. (2018), investigating unsprayed orange groves near Manaus, reported that over 50% of all predatory mites belonged to these two species, both on citrus trees and on spontaneously growing vegetation surrounding the orchards. According to the life-style classification of McMurtry et al. (2013), such dominance is typical of Type III generalist predators, which may explain their recurrence across different plant hosts in Amazonian systems.

Typhlodromips angustus was originally described from Peru, based on material from the San Martin and Amazonas administrative regions. Besides Peru, this species had been recorded in Brazil, in the state of Amazonas, by Cruz et al. (2019), in oil palm cultivation. Its dominance in guarana plantations highlights its adaptation to warm and humid climates, information that is crucial for identifying native predatory mites with potential for biological control programs in Amazonian conditions. Amblydromalus itacoatiarensis, on the other hand, is likely endemic to the Amazon (Demite et al. 2019; Brito et al. 2024, 2025) and its frequent occurrence highlights its potential relevance as a regional indicator species.

The diversity observed in this study demonstrates that guarana plantations in the Amazon harbor both widespread generalist predators and regionally restricted taxa, reinforcing their importance for documenting and understanding the Phytoseiidae fauna in this biome. Future studies should extend sampling to other guarana-producing regions and investigate seasonal dynamics and prey associations to refine the ecological interpretation of these patterns.

Acknowledgments

To Antonio C. Lofego for providing access to the phase-contrast microscopy of Acarology Laboratory, UNESP - S.J. do Rio Preto. Priscila C. Dahmer received a schorlarship (POSGRAD) from the Fundação de Amparo à Pesquisa do Estado do Amazonas (FAPEAM).

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