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Orthoptera assemblages in subtropical paddy fields: role, effect of plant phenology, and cultivation area

Published online by Cambridge University Press:  07 October 2026

Silvia Marina Albertini Open the ORCID record for Silvia Marina Albertini [Opens in a new window] ,
Mario Gabriel Ibarra-Polesel Open the ORCID record for Mario Gabriel Ibarra-Polesel [Opens in a new window] ,
Daniela Alejandra Fuentes-Rodríguez Open the ORCID record for Daniela Alejandra Fuentes-Rodríguez [Opens in a new window] ,
Martina Eugenia Pocco Open the ORCID record for Martina Eugenia Pocco [Opens in a new window] ,
Lara Milena Sabater Open the ORCID record for Lara Milena Sabater [Opens in a new window]  and
María Celeste Franceschini Open the ORCID record for María Celeste Franceschini [Opens in a new window]
Silvia Marina Albertini*
Affiliation:
Instituto de Investigaciones Geohistóricas, Consejo Nacional de Investigaciones Científicas y Técnicas, Resistencia, Chaco, Argentina Centro de Ecología Aplicada del Litoral, Consejo Nacional de Investigaciones Científicas y Técnicas, Corrientes, Argentina
Mario Gabriel Ibarra-Polesel
Affiliation:
Centro de Ecología Aplicada del Litoral, Consejo Nacional de Investigaciones Científicas y Técnicas, Corrientes, Argentina Facultad de Ciencias Exactas, Naturales y Agrimensura, Universidad Nacional del Nordeste, Corrientes, Argentina
Daniela Alejandra Fuentes-Rodríguez
Affiliation:
Facultad de Ciencias Agrarias y Forestales, Universidad Nacional de La Plata, La Plata, Argentina
Martina Eugenia Pocco
Affiliation:
Centro de Estudios Parasitológicos y de Vectores, Consejo Nacional de Investigaciones Científicas y Técnicas – Universidad Nacional de La Plata, La Plata, Argentina División Entomología, Museo de La Plata, Universidad Nacional de La Plata, La Plata, Argentina
Lara Milena Sabater
Affiliation:
Centro de Ecología Aplicada del Litoral, Consejo Nacional de Investigaciones Científicas y Técnicas, Corrientes, Argentina
María Celeste Franceschini
Affiliation:
Centro de Ecología Aplicada del Litoral, Consejo Nacional de Investigaciones Científicas y Técnicas, Corrientes, Argentina
*
Corresponding author: Silvia Marina Albertini; Email: smalbertini@gmail.com
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Abstract

Irrigated rice crops are considered artificial wetlands, with high productivity and biodiversity, especially in insect populations such as orthopterans. Although these predominantly herbivorous insects include several species of agricultural concern, their occurrence and dynamics in rice-growing systems remain insufficiently documented. This study aimed to analyse the composition and diversity of Orthoptera insects associated with subtropical paddy fields and to evaluate how rice phenology and field zone (edge vs centre) influence the structure of these assemblages. Sampling was conducted in Northeastern Argentina, the country’s main rice-producing region. In total, 14 orthopteran species belonging to four families were recorded, with Conocephalus longipes (Redtenbacher, 1891) (Tettigoniidae) as dominant, representing 89.34% of total abundance. Crop phenology significantly affected the assemblage structure: species richness and diversity peaked during the vegetative stem elongation stage, whereas overall abundance was highest during the reproductive ripening stage. In contrast, no significant differences were detected in species richness or abundance between the field edges and centres. These findings are relevant because documenting the orthopteran community present in rice systems provides insights into whether current management practices align with principles of sustainable agriculture. Moreover, the marked predominance of C. longipes suggests that this species could play an important role as a potential pest, underscoring the need for continued monitoring and targeted research to assess its impact and inform future management strategies. This study contributes to a better understanding of insect biodiversity patterns in subtropical South American paddy fields and supports efforts aimed at promoting productive yet environmentally responsible agricultural systems.

Keywords

crop stagesgrasshoppersinsect biodiversityplot variationsubtropical paddy fieldsTettigoniidae

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Type
Research Paper
Information
Bulletin of Entomological Research , First View , pp. 1 - 12
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Copyright
© The Author(s), 2026. Published by Cambridge University Press.

Introduction

Rice (Oryza sativa L.) is the crop that provides the highest caloric yield per hectare, making it one of the most important cereals worldwide (Mohapatra and Sahu, Reference Mohapatra and Sahu2022). According to the Food and Agriculture Organization of the United Nations (FAO), South America produced 25.82 million tons of rice in 2024, with Argentina ranking sixth with 1.26 million tons with an estimated income of USD 212 million in 2024 (Ministerio de Economía de la República Argentina, 2024; FAO, 2025). Production is particularly concentrated in the northeastern region of the country, with 85% of the national rice output (BCSF et al.. 2021).

Paddy fields, as agroecosystems, are distinctive for being considered as ‘artificial wetlands’ (Soldatova et al., Reference Soldatova, Dong, Li, Ivanova, Toropov, Gromyak, Dogadkin and Sun2021) and, together with natural wetlands, they play a crucial role as sources of water, productivity, and biological diversity, among other ecosystem services (Myeong and Yi, Reference Myeong and Yi2023). Because of their similarity with natural wetlands, the complexity and equilibrium of the paddy fields largely depend on their biological diversity (Propper et al., Reference Propper, Singleton, Sedlock, Smedley, Frith, Shuman-goodier and Stuart2023), allowing irrigated rice systems to reduce the negative impact of pests and weeds and to substantially enhance productivity compared with upland rice (Shekhawat et al., Reference Shekhawat, Rathore and Chauhan2020).

Orthoptera is a diverse group of insects with more than 30,000 species (Cigliano et al., Reference Cigliano, Braun, Eades and Otte2025), being particularly diverse in tropical and subtropical regions (Song, Reference Song, Foottit and Adler2018), and plays a key role in paddy fields. Due to their high levels of endemism, bioindicator quality (Stefanidis et al., Reference Stefanidis, Zografou, Tzortzakaki and Kati2023), and large body size, they could be useful for assessing social–environmental conditions in wetlands and agroecosystems. Functionally, orthopterans are essential in trophic dynamics; being predominantly herbivores, they can act as major crop pests or contribute to weed control, while serving as prey for numerous predators and parasitoids (Giuliano et al., Reference Giuliano, Cardarelli and Bogliani2018; Meena and Meena, Reference Meena and Meena2021; Srygley, Reference Srygley, Eigenbrode and Rashad2023). Most available information on Orthoptera in paddy fields comes from Asia, where 90% of the global rice production occurs (Chitra et al., Reference Chitra, Soundararajan and Gunathilagaraj2000; Sultana and Wagan, Reference Sultana and Wagan2009; Ane and Hussain, Reference Ane and Hussain2016; Gyawali et al., Reference Gyawali, Regmi, Pudasaini and Acharya2019; Takeuchi and Irie, Reference Takeuchi and Irie2019; Cuong et al., Reference Cuong, Souvannalath, Ha, Lien, Tam, Hoai and Dat2022). The reports include several genera reported as pests (Sultana and Wagan, Reference Sultana and Wagan2009; Ane and Hussain, Reference Ane and Hussain2016), as well as species that act as predators of pest eggs (Chitra et al., Reference Chitra, Soundararajan and Gunathilagaraj2000). For example, several studies conducted in Asia have reported species of the genus Conocephalus (Orthoptera: Tettigoniidae) as potential rice pests and predators of pest eggs, highlighting their potentially dual role in rice agroecosystems (Corrales Castillo et al., Reference Corrales Castillo, Moya, Martínez, Arrieta and Herrera2017; Hegazy et al., Reference Hegazy, Hendawy, Mesbah and Salem2021; Suroto et al., Reference Suroto, Istiqomah and Syarifah2021; Tandon and Srivastava, Reference Tandon and Srivastava2021). We also found Conocephalus longipes (Orthoptera: Tettigoniidae) in paddy fields in northeastern Argentina; however, its ecological role is still unclear. In the Americas, several Orthoptera species have also been reported in association with rice agroecosystems, including species of different genera (Reyna et al., Reference Reyna, Trabanino, Avedillo, Pitty and Rueda1994; Le-Gall et al., Reference Le-Gall, Overson and Cease2019) and Dichroplus elongatus Giglio-Tos in Brazil and Argentina (Lange and Cigliano, Reference Lange, Cigliano, Lecoq and Zhang2019; Mariottini et al., Reference Mariottini, Mancini, Falasco, Wysiecki and Lange2023).

In rice plots, irrigation water is distributed through channels (primary, secondary, tertiary, and field channels) (Kraemer et al., Reference Kraemer, Moulin, Marín, Kruger and Herber2024) originating from rivers or reservoirs, which promotes the establishment of aquatic macrophytes, especially along the irrigation channels and field margins. Paddy fields are also commonly surrounded by native forest and grasslands, increasing the biodiversity at the crop margins, a pattern known as ‘edge effect’ (Pérez-Méndez et al., Reference Pérez-Méndez, Alcaraz and Catala-Forner2025). These surrounding habitats act as refuges for various arthropod species, including rice pests that overwinter in these habitats (Fuentes-Rodríguez et al., Reference Fuentes-Rodríguez, Franceschini, Gervazoni, López, Sosa and Kruger2020), while the greater plant diversity also supports natural enemies and enhances biological control through habitat management (Berensztein et al., Reference Berensztein, Ricci, Polack and Marasas2017; Zou et al., Reference Zou, De Kraker, Bianchi, Xiao, Huang, Deng, Hou and Van der Werf2020). Furthermore, alteration between flooded and dry phases in irrigated rice systems provides diverse habitat conditions and ecological niches that could be colonised by different animal taxa, including insects and other invertebrates. Altogether, these features make paddy fields more similar to natural wetlands than to other kinds of terrestrial crops such as soybean, cotton, sunflower, among others (Yoon, Reference Yoon2009; Maltchik et al., Reference Maltchik, Rolon, Stenert, Machado and Rocha2011; Mowjood et al., Reference Mowjood, Jinadasa and Basnayake2020).

In this context, although several studies have evaluated how insect communities vary with rice crop phenology and field area (Krinski and Foerster, Reference Krinski and Foerster2017; Costa et al., Reference Costa, Seidel, Pazini, da Silva, da Silva, da Silva Martins and Barrigossi2019; Fuentes-Rodríguez et al., Reference Fuentes-Rodríguez, Dellapé, Gervazoni, Giese, Prieto, Sosa and Franceschini2025), they have mainly focused on insect groups that include main rice pests, such as hemipterans and lepidopterans. However, much less attention has been given to orthopteran assemblages. Addressing these aspects in Orthoptera is relevant for understanding the dynamics of current and potential pest species, as well as for assessing their potential use in biodiversity monitoring and as indicators of good agricultural practices.

There is limited knowledge about the functional role, abundance, diversity, and community composition of Orthoptera in tropical and subtropical paddy fields, as well as how these parameters vary temporally with crop phenology and spatially across different areas of the field. Against this background, especially, production focuses on the implementation of good agricultural practices and integrated pest management strategies that promote both economic and environmental sustainability, the study of orthopteran assemblages, especially their functional roles (Gangwere, Reference Gangwere1961), trophic level, and specificity of host degrees, would provide valuable information to determine if orthopterans are relevant as rice pests or as biocontrol agents against other rice pest species.

The aim of this study was to evaluate Orthoptera assemblages in relation to rice crop phenology and spatial variability between field edges and centres, to assess the role these insects play in rice agroecosystems. Our hypotheses were as follows: (a) species richness, abundance, diversity, and composition of Orthoptera assemblages associated with subtropical rice systems vary with phenological stages of rice crops, reaching higher values during reproductive stages due to increased structural complexity which provides more microhabitats for orthopterans; (b) these parameters will be higher at field edges than at centres because of the greater plant heterogeneity at the margins of the paddy fields.

Material and methods

Study area and cultivation system: Sampling was conducted in 10 paddy fields located in Northeastern Argentina, in Corrientes province (fig. 1). Sampling was carried out over the rice production cycle, between November 2017 and April 2018. According to the Köppen–Geiger system, the region has humid subtropical weather (Cfa) (CRU and GPCC, 2022), with mean annual temperatures of 21.7°C and an average annual precipitation of 1289 mm.

Figure 1.

Subtropical paddy fields in Northeastern Argentina where the samplings of Orthoptera were made: Site 1 (27°25′07.7″S, 57°38′05.8″ W), Site 2 (27°23′16.0″ S, 57°41′00.9″ W), Site 3 (27°28′53.7″ S, 57°27′58.0″ W), Site 4 (27°49′44.8″ S, 58°46′20.2″ W), Site 5 (27°49′51.5″ S, 58°48′10.4″ W), Site 6 (27°40′21.1″ S, 58°48′09.2″ W), Site 7 (27°29′17.2″ S, 57°23′55.7″ W), Site 8 (27°25′53.9″ S, 57°22′53.3″ W), Site 9 (27°17′30.9″ S, 58°17′25.0″ W), and Site 10 (27°18′40.0″ S, 58°18′30.9″ W).

Sampling sites in Northeastern Argentina near the Paraguay and Parana Rivers, with labeled provinces.

The rice crop presents different phenological stages characterised by changes in growth and development. In this research, four stages were analysed: tillering (a rapid growth period with high nutrient requirement; vegetative phenology (VP)), stem elongation (high growth rates and panicle differentiation; VP), flowering (panicle emergence and anthesis; reproductive phenology (RP)) and ripening (grain filling and maturation; RP) (Kruger and Burdyn, Reference Kruger and Burdyn2015; Qin et al., Reference Qin, Hu, Yuan, Liu, Wang, Liu, Guo and Song2023) (fig. 2).

Figure 2.

Spatial variation of paddy fields in Northeastern Argentina: (1a) native forest, (1b) native grassland, (2) main irrigation channel, (3) crop edge, (4) crop centre, and (5) secondary irrigation channel. Each white dot marks a sampling point, with 10 m between adjacent points. Temporal variation: vegetative stages of rice development (September–December) and reproductive stages (January–March).

Scientific illustration with schematic and aerial photo showing spatial and temporal variation in paddy fields, labeled with numbered sites. See long description.
Figure 2 Long description

The illustration consists of two panels. The schematic panel at the top is labeled, Spatial variation and depicts different areas: native forest labeled 1a, native grassland labeled 1b, main irrigation channel labeled 2, crop edge labeled 3, crop center labeled 4 and secondary irrigation channel labeled 5. Above this, the schematic shows, Temporal variation, with vegetative stages from September to December and reproductive stages from January to March. The aerial-photo panel below shows the same labeled areas from above, with white dots marking sampling points in the crop edge and crop center. The layout is functional, with clear callouts and panel headings to convey scientific data on paddy field variation.

In contrast, the sampled fields presented irrigation channels that supported aquatic macrophytes and fauna, especially birds and invertebrates (Maltchik et al., Reference Maltchik, Rolon, Stenert, Machado and Rocha2011). In the sampled paddy fields, the crop starts under dry conditions, and irrigation begins after 3 weeks of seedling emergence. The main channels connect with the Paraná River and Paraguay River, from which irrigation water was extracted. Water supply is interrupted 20 days after the flowering stage starts, to allow harvest under dry soil conditions (Kraemer et al., Reference Kraemer, Moulin, Marín, Kruger and Herber2024). Additionally, field centres are characterised for the presence of O. sativa L. almost exclusively, whereas field edges also contained other floating and emergent aquatic plants (e.g. Pontederia azurea Sw, Sagittaria montevidensis Cham & Schltdl, Salvinia sp., Nymphoides indica (L.) Kuntze, and Ludwigia sp.) as well as native grass (Paspalum sp., Cyperus sp., Elionorus sp., and Eryngium sp., among others) (fig. 2).

Sampling procedure

Ten paddy fields were sampled through all the phenological stages of the rice plant. The paddy fields were separated by more than 2 km to ensure sample independence. Insects were captured using an entomological net of 30 cm in diameter and 50 cm in depth.

To analyse temporal variation of Orthoptera assemblages, each paddy field was sampled once during each of the four phenological stages of the annual rice-growing cycle. Sampling dates were determined according to the phenological development of the crop rather than at fixed time intervals. To assess the spatial variation at the local scale, two transects were established within each rice plot: one along the edge of the plot and another in the centre. The sampling unit was a sampling plot measuring 5 m × 1 m. Within each paddy field, two transects were established, one along the field edge and another in the field centre. Each transect contained five sampling plots separated by 10 m, resulting in 10 sampling plots per paddy field at each phenological stage. At each sampling plot, three consecutive net sweeps were performed over the vegetation every two steps, totalling nine sweeps per sampling plot. Thus, each rice field contributed 10 sampling plots × 4 phenological stages = 40 samples, and the overall sampling effort comprised 400 samples (10 rice fields × 4 phenological stages × 10 sampling plots). To enhance data randomisation, the location of the first sampling plot in each transect was determined using randomly generated numbers.

Orthopterans collected were separated and preserved in 70% ethanol. In the laboratory, specimens were mounted, labelled, and subsequently identified using specialised taxonomic keys and databases (Carbonell et al., Reference Carbonell, Cigliano and Lange2025; Cigliano et al., Reference Cigliano, Braun, Eades and Otte2025). Material was deposited in the Arthropod Collection (CARTROUNNE) of Facultad de Ciencias Exactas, Naturales y Agrimensura, Universidad Nacional del Nordeste (FaCENA-UNNE).

For the assemblage analysis, the richness, abundance, diversity, and composition were estimated considering phenological stages and field zones (edge-centre). For each Orthoptera species sampled, information on the ecological role and feeding habit (polyphagous, oligophagous, or monophagous) was reviewed from the available bibliographic sources. Species were also categorised as potential rice pests or potential natural enemies of insect pests based on published information.

Statistical analysis

To determine if differences exist in Orthoptera assemblage attributes (richness, abundance, diversity, and composition) among rice phenological stages (tillering, stem elongation, flowering, and ripening), the non-parametric Friedman test for repeated measures was performed followed by post hoc pairwise comparisons, using InfoStat software (Di Rienzo et al., Reference Di Rienzo, Casanoves, Balzarini, Tablada, Gonzales and Robledo2017). To evaluate differences in Orthoptera abundance between life stages (nymphs and adults) and crop phenological phases, abundance data were grouped into vegetative (tillering and stem elongation) and reproductive (flowering and ripening) phases, and analysed using a two-way analysis of variance (ANOVA), with phenological phase and life stage as fixed factors, including their interaction. Diversity was evaluated using Hill numbers, or effective species numbers (Jost, Reference Jost2006), where the exponent q determines the sensitivity index to relative abundances; that is, the extent to which common or rare species (those with ≤9 individuals) influence the diversity measure. For this analysis, first-order diversity (1D; the exponential of Shannon entropy) and second-order diversity (2D; the inverse of Simpson’s index) were calculated. Diversity indices were calculated using the PAST 4.03 software (Hammer et al., Reference Hammer, Harper and Ryan2020).

Differences in orthopteran richness and abundance between the edge and centre of the crop were analysed with Kruskal–Wallis test. To evaluate if differences exist in the composition of orthopteran assemblages between the edge and centre of the crop, an analysis of similarity (ANOSIM) was performed. The similarity among species composition in the different crop areas was calculated with the incidence-based complementarity index and the abundance-based Bray–Curtis index. To analyse the influence of the different rice plot areas on orthopteran abundance, comparisons were made using Whittaker rank–abundance curves (Magurran, Reference Magurran2004). To examine the spatial distribution patterns of species abundances that constitute the grasshopper assemblages in the two areas, a non-metric multidimensional scaling (NMDS) ordination was conducted. All the analyses mentioned were calculated with the PAST 4.03 software (Hammer et al., Reference Hammer, Harper and Ryan2020). The significance level considered was p = 0.05.

Results

A total of 638 orthopteran specimens were collected across all paddy fields and phenological stages, belonging to 14 species, 14 genera, and four families (table 1). The family with the greatest species richness was Acrididae (S = 10; n = 59), while the most abundant was Tettigoniidae (S = 2; n = 576), whereas Tetrigidae and Romaleidae were each represented by a single species. The assemblage was strongly dominated by Conocephalus longipes (Redtenbacher) (Tettigoniidae), which represented 89.34% of the total abundance (n = 570), followed by Aleuas vitticollis Stål (3.1%; n = 20).

Table 1.

Abundance of Orthoptera species and community descriptors across crop phenological stages and spatial zones in subtropical paddy fields of Northeastern Argentina

Conocephalus longipes had the highest counts, rising from 41 at tillering to 237 at ripening, with totals increasing from 43 to 258 across stages. See long description.
Table 1 Long description

Counts are given for Orthoptera species by family across phenological stages, tillering, stem elongation, flowering, and ripening, plus totals for edge and center crop areas. Conocephalus longipes is listed at 41 in tillering, 71 in stem elongation, 221 in flowering, and 237 in ripening; by area it is 268 at edge and 302 at center. Total abundance by stage is 43 in tillering, 90 in stem elongation, 247 in flowering, and 258 in ripening; by area totals are 306 at edge and 332 at center. Total richness by stage is 3 in tillering, 13 in stem elongation, 7 in flowering, and 8 in ripening; by area richness is 12 at edge and 10 at center. First-order diversity 1D is 1.2 in tillering, 2.7 in stem elongation, 1.6 in flowering, and 1.5 in ripening; by area it is 1.9 at edge and 1.6 at center. Second-order diversity 2D is 1.1 in tillering, 1.6 in stem elongation, 1.2 in flowering, and 1.2 in ripening; by area it is 1.3 at edge and 1.2 at center. Other species with nonblank stage values include Aleuas vitticollis at 3 in stem elongation, 10 in flowering, and 7 in ripening; Belosacris coccineipes at 1 in tillering, 2 in stem elongation, 7 in flowering, and 5 in ripening; and Metaleptea adspersa at 5 in stem elongation, 3 in flowering, and 1 in ripening.

Notes: The upper section presents species-level abundances grouped by family; the lower section presents summary community metrics (total abundance, species richness, 1D and 2D). Phenological stages: tillering (Til), stem elongation (SE), flowering (Flo), and ripening (Rip). Crop areas: edge and centre.

Effects of rice crop phenology on the Orthoptera assemblages: richness, abundance, diversity, and composition

In relation to Orthoptera richness, no significant differences among phenological stages were found (Friedman test, T 2 = 2.19, p = 0.1121). The highest values were recorded during stem elongation (vegetative stage; S = 13) and lower values in the tillering stage (vegetative stage; S = 3) (fig. 3).

Figure 3.

Mean richness (log10) of Orthoptera in subtropical paddy fields of Northeastern Argentina in relation to vegetative (Til = tillering; SE = Stem elongation) and reproductive phenological stages (Flo = flowering; Rip = ripening).

A bar graph showing mean richness of Orthoptera across phenological stages.

Concerning the abundance, most of the specimens were recorded during reproductive stages (n = 505; 79.15%), with significant differences among the four phenological stages (Friedman test, T 2 = 3.23, p = 0.0379) (fig. 4A). Abundance was significantly lower during tillering than during flowering and ripening, whereas stem elongation did not differ significantly from either vegetative or reproductive stages. Considering life stages, nymphs predominated over adults throughout the crop cycle (ANOVA, F 1.27 = 6.65, p = 0.0157), with total abundance being significantly higher during the reproductive stage (ANOVA, F 1.27 = 12.38, p = 0.0016) (fig. 4B). Nevertheless, the interactions between phenological stage and life stage were not statistically significant (ANOVA, F 1.27 = 2.95, p = 0.0971). Regarding diversity, the highest values of this parameter were found during the stem elongation vegetative stage (1D = 2.7; 2D = 1.6), whereas the lower values were observed in the tillering vegetative stage (1D = 1.2; 2D = 1.1) (table 1).

Figure 4.

(A) Mean abundance (log10) of the Orthoptera assemblage observed in subtropical paddy fields at Northeastern Argentina in relation to vegetative (Til = tillering; SE = stem elongation) and reproductive phenological stages (Flo = flowering; Rip = ripening). (B) Mean abundance (log10) of adult and nymphal Orthoptera comparing vegetative (Veg) and reproductive (Rep) phenology of rice crops in Northeastern Argentina. Significant differences are indicated by different letters above the bars (post hoc test).

Two bar graphs showing mean abundance of Orthoptera across phenological stages and comparing nymphs and adults in vegetative versus reproductive stages. See long description.
Figure 4 Long description

A showing a bar graph with the vertical axis label, Mean abundance of Orthoptera, with units not specified and tick labels 2, 10, 18, 26, 34. The horizontal axis label is, Phenological stages, with category labels Til, SE, Flo, Rip. A p value annotation reads, p equals 0.0379. Four bars are shown with error bars. Approximate bar heights are Til about 3, SE about 9, Flo about 22 and Rip about 23. Letters above the bars read A above Til, AB above SE, B above Flo and B above Rip. B showing a grouped bar graph with the vertical axis label, Mean abundance of Orthoptera, with units not specified and tick labels 3, 16, 29, 42, 55. The horizontal axis label is, Phenological stages, with category labels Veg and Rep. A legend lists Nymphs and Adults, shown as two different bar styles. A p value annotation reads, p equals 0.0016. At Veg, the Nymphs bar is about 11 and the Adults bar is about 4, with error bars. At Rep, the Nymphs bar is about 43 and the Adults bar is about 6, with error bars. Letters above the grouped bars read A above Veg and B above Rep and the letter A appears above the Adults bar at Veg and above the Adults bar at Rep.

With respect to the assemblage composition, the vegetative stem elongation stage presented 13 of the recorded species, being Adimantus ornatissimus (Burmeister), Cylindrotettix santarosae (Roberts), Paulinia acuminata (De Geer), Stenacris xanthochlora (Marschall), and Tetrigidae sp. exclusive to this stage. In contrast, the reproductive ripening stage presented eight of the recorded species, followed by the flowering stage (S = 7) with Leptysma argentina Bruner as the exclusive species in this stage. The vegetative tillering stage recorded only three of the species. In addition to this, the species present in all the phenological stages were C. longipes (Redtenbacher) (Tettigoniidae) and Belosacris coccineipes (Bruner) (Acrididae).

Effects of field zone (edge vs. centre) on Orthoptera assemblage: richness, abundance, diversity, and composition

The crop edges tended to present a higher richness (S = 12) than the centre (S = 10) of the paddy field; however, these differences were not significant (Kruskal–Wallis test, H = 1.195; p = 0.1986) (table 1).

In relation to the abundance, no significant differences were found between edges and centres (Kruskal–Wallis test, H = 0.12; p = 0.74), with similar proportions of individuals in both crop areas. The Whittaker (ranking-abundance) curves showed a very similar distribution of the abundance in the orthopteran species present in the edge and centre of the paddy fields (fig. 5). Both curves revealed similar patterns, with a strong dominance by C. longipes and many rare species. This pattern was supported by the Bray–Curtis index (based on abundance), indicating a high similarity between the two field areas (91%). Diversity was slightly higher in the field-edge zone (1D = 1.9; 2D = 1.3) compared to the centre (1D = 1.6; 2D = 1.2) (table 1).

Figure 5.

Ranking-abundance curves in two areas of the plots (edge and centre) of paddy fields in Northeastern Argentina. References: Clong (Conocephalus longipes); Avit (Aleuas vitticolis); Bcoc (Belosacris coccineipes); Dexi (Dichroplus exilis); Madsp (Metaleptea adspersa); Etrun (Eucaulopsis truncata); Larg (Leptysma argentina); Tgrac (Tucayaca gracilis), Cang (Coryacris angustipennis); Cysant (Cylindrotettix santarosae); Pacum (Paulinia acuminata); Sxant (Stenacris xanthochlora); Tetri sp. 1 (Tetrigidae sp.); and Aorn (Adimantus ornatissimus).

Ranking-abundance curves compare species abundance at field edges and centers. See long description.
Figure 5 Long description

Ranking-abundance curves for orthopteran species in two areas: field edge and field center. The vertical axis represents abundance in logarithmic scale, labeled as log superscript 10. The horizontal axis is divided into two sections: Field Edge and Field Center. Each section displays species names along the curve, indicating their abundance. In both sections, the species Clong is dominant, with the highest abundance. Other species such as Avit, Bcoc, Dexi, Madsp, Etrun, Larg, Tgrac, Cang, Cysant, Pacum, Sxant, Tetri sp. 1 and Aorn are listed with decreasing abundance. The curves show similar patterns in both areas, with Clong having the highest abundance and other species showing lower values.

About assemblage composition, most of the species were found in both crop areas (edge and centre; S = 8; 57.14%), while four species were exclusive to edges (S. xanthochlora, C. santarosae, P. acuminata, and Tetrigidae sp.; 28.57%) and two to centres (A. ornatissimus, L. argentina; 14.28%) (fig. 6). Turnover values between crop areas were moderate (C = 0.4) according to the complementarity index based on species incidence. The NMDS analysis showed that the edge and centre areas of the rice crop formed two groups that were partially overlapping (fig. 7), supported by the ANOSIM, which detected no significant differences in the orthopteran composition between both areas (R = −0.086; p = 0.7657).

Figure 6.

Venn diagram with the Orthoptera species exclusive to the edges and the centre of the paddy fields, as well as those occurring in both zones. C represents the species turnover value between the two field zones.

A 2-set Venn diagram showing Orthoptera species distribution between field edges and centers.
Figure 7.

Non-metric multidimensional scaling (NMDS) analysis of the orthopteran assemblage in subtropical paddy fields of Northeastern Argentina. Circles represent the replicate sites located in the field-edge areas, whereas squares represent the replicates from the field-centre areas.

A scatter plot showing NMDS analysis of orthopteran assemblage with Edge and Center sites represented by circles and squares.

Role of the Orthoptera assemblages in subtropical rice paddies

The analysis of the information reviewed about feeding habits of the collected orthopterans showed that 11 species were classified as herbivorous, whereas 1 was omnivorous. Additionally, among the herbivorous species, all were generalists, with seven graminivorous species. No oligophagous herbivore species were recorded (table 2).

Table 2.

Trophic level and feeding habits of Orthoptera based on the available literature in subtropical paddy fields of Northeastern Argentina and role as potential pesta of rice, potential natural enemies of pest insectsb, and bioindicatorsc

Conocephalus longipes had the highest total abundance at 570, far exceeding all other Orthoptera species listed. See long description.
Table 2 Long description

Columns list Family, Species, Total Abundance, Trophic level and feeding habits, and Reference for Orthoptera species. Families shown are Acrididae, Romaleidae, Tetrigidae, and Tettigoniidae. Total abundance values range from 1 to 570, with Conocephalus longipes in Tettigoniidae at 570 and described as omnivorous. In Acrididae, abundances are 20 for Aleuas vitticollis, 15 for Belosacris coccineipes, 9 for Metaleptea adspersa, 6 for Dichroplus exilis, 3 for Tucayaca gracilis, 2 for Leptysma argentina, and 1 each for Stenacris xanthochlora, Cylindrotettix santarosae, Paulinia acuminata, and Adimantus ornatissimus. Romaleidae includes Coryacris angustipennis with abundance 2 and feeding listed as herbivorous, forbivorous or polyphagous. Tetrigidae includes Tetrigidae sp. with abundance 1 and feeding shown as a single hyphen. Tettigoniidae also includes Eucaulopsis truncata with abundance 6 and trophic level and feeding habits listed as ND.

Notes: ND, no data available.

Discussion

This research characterises Orthoptera assemblages and their role in subtropical paddy fields, evaluating their temporal (rice phenology) and spatial (field edge vs centre) variation. Although previous studies about orthopteran assemblages were conducted in paddy fields of Asia and Europe (Giuliano and Bogliani, Reference Giuliano and Bogliani2018; Jauharlina et al., Reference Jauharlina, Hasnah and Taufik2019; Abrori and Leksono, Reference Abrori and Leksono2021), their results cannot be extrapolated to South America because of the differences in orthopteran species, climate, and management practices. Furthermore, since orthopteran collection demands specialised sampling techniques, the value of this study is reinforced, given its simultaneous assessment of the effects of crop phenology and field zone variation on the assemblages.

The dominance of the families Tettigoniidae and Acrididae in this study is consistent with reports in paddy fields of Central and South America (Reyna et al., Reference Reyna, Trabanino, Avedillo, Pitty and Rueda1994; Fritz et al., Reference Fritz, Heinrichs, Machado, Andreis, Pandolfo, de Salles, de Oliveira and Fiuza2011; Cervantes and Huacuja, Reference Cervantes and Huacuja2015; Acosta et al., Reference Acosta, Jahnke, Redaelli and Pires2017; Ghiglione et al., Reference Ghiglione, Zumoffen, Dalmazzo, Strasser and Attademo2021; Menegon and Jahnke, Reference Menegon and Jahnke2021), Asia and Europe (Fathima et al., Reference Fathima, Pirya, Meeran, Arivoli and Tennyson2021; Menegon and Jahnke, Reference Menegon and Jahnke2021; Mathew et al., Reference Mathew, Joseph and Cyril2022), suggesting that these families are well adapted to rice agroecosystems, regardless management practices, climate, and regions. Additionally, this research provides five species recorded for the first time in the study area (A. ornatissimus, A. vitticollis, B. coccineipes, L. argentina, and T. gracilis), contributing to future checklists, taxonomic revisions of these species, and biodiversity inventories.

Regarding our hypotheses, the results obtained in this study support the first hypothesis partially, indicating that Orthoptera are sensitive to phenological changes in paddy fields. Although we predicted that the reproductive stages would exhibit the highest species richness, abundance, and diversity, only abundance followed this pattern. In contrast, species richness and diversity reached their highest values during the stem elongation stage, a vegetative stage of crop development. On the other hand, the second hypothesis was not supported, as none of the evaluated parameters (species richness, abundance, diversity, and composition) showed significant differences between the edge and centre zones of the paddy fields. Although species richness and diversity tended to be slightly higher at the field edges, these differences were not statistically significant.

Rice phenological stages provide orthopterans with different food sources, refugee, and habitat, due to changes in nutritional quality and architecture of plants, as well as the presence of other insects (e.g. Pentatomidae, Cicadellidae, Libellulidae, Formicidae, among others; Paradell et al., Reference Paradell, Maciá, Asborno, Catalano, Brentassi, Varela, Dellapé, Defea, Foieri and de Remes Lenicov2014; Abidin et al., Reference Abidin, Leksono, Yanuwiadi and Purnomo2020; Fuentes-Rodríguez et al., Reference Fuentes-Rodríguez, Dellapé, Gervazoni, Giese, Prieto, Sosa and Franceschini2025) that coexist and compete with Orthoptera for resources. Structural changes in rice plants include stem thickening, flag leaf and panicle emergence, anthesis, grain formation, and mobilisation of sugars (Lai and Lin, Reference Lai and Lin2021), which could explain the variations observed in richness, abundance, diversity, and composition in our study. Similar patterns have been documented in other rice systems (Saavedra de Castro and Fernández-Herrera, Reference Saavedra de Castro and Fernández-Herrera2019) and crops (Mathew et al., Reference Mathew, Joseph and Cyril2022; Rodbell et al., Reference Rodbell, Kaur and Schell2025).

Our results show a higher richness of orthopteran species in vegetative stages, which is supported with previous results reported by Machado and Garcia (Reference Machado and Garcia2010) in Brazilian paddy fields. This pattern could be related to a higher succulence in the leaves during vegetative stages, making them potentially more attractive as a food source for orthopterans (Gangurde, Reference Gangurde2007). Contrary to our results, however, other studies have documented higher richness of Orthoptera species in reproductive stages of rice crops (Edirisinghe and Bambaradeniya, Reference Edirisinghe and Bambaradeniya2006; Roy et al., Reference Roy, Kabir, Akter, Nayeem, Alam, Hasan, Bari and Sannal2024).

Although Orthoptera assemblages varied across phenological stages, no spatial differences (edge vs centre) were detected in richness or abundance, whereas diversity differed between both zones. Composition did not vary either. Our results show similar Orthoptera species richness at the field edge and centre, in contrast with findings reported for other invertebrate communities in paddy fields (Dominik et al., Reference Dominik, Seppelt, Horgan, Settele and Václavík2018) and other crop systems (Molina et al., Reference Molina, Poggio and Ghersa2016; Pérez-Mendez et al., Reference Pérez-Mendez, Pla, Tomás, Bartomeu, Ferré and Catala-Forner2020; Ghiglione et al., Reference Ghiglione, Zumoffen, Dalmazzo, Strasser and Attademo2021), where richness is higher at field edges. These patterns are usually attributed to higher plant diversity and greater resource availability at field edges, which are adjacent to irrigation channels and native vegetation (Fuentes-Rodríguez et al., Reference Fuentes-Rodríguez, Dellapé, Gervazoni, Giese, Prieto, Sosa and Franceschini2025). However, our results suggest that Orthoptera assemblages in paddy fields respond more strongly to plant phenology than to edge effects. This may be explained by simpler plant diversity at field edges, due to disturbances such as grazing, burning, and pesticide application (Sáez et al., Reference Sáez, Sabatino and Aizen2014). A study by Pérez-Méndez et al. (Reference Pérez-Méndez, Alcaraz and Catala-Forner2025) suggests that the most effective way to support beneficial insects may be the restoration of native vegetation along the edges of paddy fields, even though passive restoration, as it provides refuge and food resources for the natural enemies of rice pests, including predators and parasitoids.

In relation to the recorded orthopteran species, the dominance of polyphagous herbivorous species, particularly graminivores, together with the dominance of C. longipes, whose genus has been reported as an occasional rice pest (Chitra et al., Reference Chitra, Soundararajan and Gunathilagaraj2000; Ane and Hussain, Reference Ane and Hussain2016; Corrales Castillo et al., Reference Corrales Castillo, Moya, Martínez, Arrieta and Herrera2017; Suroto et al., Reference Suroto, Istiqomah and Syarifah2021), should be considered in monitoring programs of potential pests that may emerge once current pests are controlled or under changes in environmental conditions. For example, species such as Melanoplus sp., Sphenarium mexicanum Saussure, 1859, Sphenarium purpurascens Charpentier, 1845, Brachystola magna (Girard, 1854) and Romalea eques Burmeister, 1838, are usually regarded as secondary or economically insignificant pests, but after intense rainfall or severe drought they may become endemic pests, causing substantial economic losses (Mariño-Pérez et al., Reference Mariño-Pérez, Fontana and Buzzetti2011).

Irrigated paddy fields are artificial wetlands that harbour species commonly found in natural wetlands. In the orthopteran assemblages surveyed, several species (M. adspersa, T. gracilis, P. acuminata, C. angustipennis) and genera (Aleuas, Dichroplus, Leptysma, Conocephalus) have also been recorded in natural wetlands (Capello et al., Reference Capello, Marchese and De Wysiecki2013). Several of these species exhibit adaptations to aquatic environments, including endophytic or epiphytic oviposition, broadened hind tibiae for swimming, and a fusiform body shape for submersion (Nunes-Gutjahr and de Souza Braga, Reference Nunes-Gutjahr and de Souza Braga2018). Our results confirm that paddy fields support wetland-characteristic Orthoptera biodiversity, consistent with previous studies focused on different taxonomic groups (Maltchik et al., Reference Maltchik, Rolon, Stenert, Machado and Rocha2011; Choe et al., Reference Choe, Cho, Han, Kim, Choi, Bang, Eo, Na and Kim2016; Paulino et al., Reference Paulino, Granadeiro, Matos and Catry2025). Considering the use of orthopterans as bioindicators (Tan et al., Reference Tan, Yeo and Hwang2017; Fumy et al., Reference Fumy, Löffler, Samways and Fartmann2020; Abd Rahim et al., Reference Abd Rahim, Muhammad and Abdullah2024), and the overlap with natural wetlands species, Orthoptera may represent a promising group for assessing the ecological effects of paddy field management. However, future studies comparing different management practices are needed to evaluate their potential as indicators of sustainable management.

Regarding the dominance of C. longipes, its high abundance may have influenced Orthoptera richness and diversity in paddy fields. This pattern could reflect C. longipes’ greater adaptation to rice agroecosystems, as well as a superior competitive ability for available resources relative to other Orthoptera species (Avolio et al., Reference Avolio, Forrestel, Chang, La Pierre, Burghardt and Smith2019). On the other hand, more studies are needed to clarify the ecological role and feeding habits of Conocephalus species in rice agroecosystems. Some studies affirm that some Conocephalus species act as predators of the eggs of rice pests, particularly hemipterans and lepidopterans, and may therefore play a role as biocontrol agents in rice crops (Ito et al., Reference Ito, Kin and Chang1995; Oda and Ishii, Reference Oda and Ishii2001; Chitra et al., Reference Chitra, Gunathilagaraj and Soundararajan2002; Gangurde, Reference Gangurde2007; Ane and Hussain, Reference Ane and Hussain2016; Acosta et al., Reference Acosta, Jahnke, Redaelli and Pires2017; Borkakati et al., Reference Borkakati, Saikia and Buragohain2018; Iqbal, Reference Iqbal, Riaz and Zaman2020; Hegazy et al., Reference Hegazy, Hendawy, Mesbah and Salem2021; Tandon and Srivastava, Reference Tandon and Srivastava2021). Other studies indicate that Conocephalus can feed on pollen, panicles, and rice leaves, being particularly abundant during the reproductive stages of the crop (Reyna et al., Reference Reyna, Trabanino, Avedillo, Pitty and Rueda1994; Ito et al., Reference Ito, Kin and Chang1995; Chitra et al., Reference Chitra, Gunathilagaraj and Soundararajan2002), including several species that are considered pests of crops such as rice (Chitra et al., Reference Chitra, Soundararajan and Gunathilagaraj2000; Oda and Ishii, Reference Oda and Ishii2001; Ane and Hussain, Reference Ane and Hussain2016; Corrales Castillo et al., Reference Corrales Castillo, Moya, Martínez, Arrieta and Herrera2017; Suroto et al., Reference Suroto, Istiqomah and Syarifah2021). This highlights the plasticity of the genus and the need for more ecological and taxonomic studies to assess its potential impact in the region.

Finally, our results contribute to understanding Orthoptera biodiversity in subtropical paddy fields by demonstrating that assemblages are primarily structured by crop phenology rather than plot zones. The overlap of species with natural wetlands reinforces the value of these assemblages as bioindicators of ecological conditions and highlights their potential use in sustainable management strategies in rice agroecosystems.

Acknowledgements

The authors used an AI-based language tool to assist with translation and English editing. All content was reviewed and approved by the authors. The authors extend their gratitude to Technician Pedro Quaranta for preparing the illustrations and rice producers from the study areas for their collaboration.

Author contributions

S.M.A. curated the data, performed statistical analyses, prepared figures, and wrote the original draft of the manuscript. M.G.I.P., D.A.F.R., and L.M.S. contributed to figure preparation, supervision, and manuscript review and editing. M.E.P. carried out taxonomic identification and contributed to manuscript review and editing. M.C.F. was responsible for funding acquisition, project administration, supervision, taxonomic identification, and manuscript review and editing.

Ethics and approvals

All authors adhered to the ethical policies of CONICET-UNNE.

Financial support

This study was funded by Argentina’s National Scientific and Technical Research Council (CONICET: PIP 2014-2016 no. 0331 and PIP 11220130100533) and Argentina’s National Agency for the Promotion of Science and Technology (ANPCyT: PICT 1910-2015).

Competing interests

The authors declare that no conflicts of interest exist.

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Figure 0

Figure 1. Subtropical paddy fields in Northeastern Argentina where the samplings of Orthoptera were made: Site 1 (27°25′07.7″S, 57°38′05.8″ W), Site 2 (27°23′16.0″ S, 57°41′00.9″ W), Site 3 (27°28′53.7″ S, 57°27′58.0″ W), Site 4 (27°49′44.8″ S, 58°46′20.2″ W), Site 5 (27°49′51.5″ S, 58°48′10.4″ W), Site 6 (27°40′21.1″ S, 58°48′09.2″ W), Site 7 (27°29′17.2″ S, 57°23′55.7″ W), Site 8 (27°25′53.9″ S, 57°22′53.3″ W), Site 9 (27°17′30.9″ S, 58°17′25.0″ W), and Site 10 (27°18′40.0″ S, 58°18′30.9″ W).

Figure 1

Figure 2. Spatial variation of paddy fields in Northeastern Argentina: (1a) native forest, (1b) native grassland, (2) main irrigation channel, (3) crop edge, (4) crop centre, and (5) secondary irrigation channel. Each white dot marks a sampling point, with 10 m between adjacent points. Temporal variation: vegetative stages of rice development (September–December) and reproductive stages (January–March).Figure 2 long description.

Figure 2

Table 1. Abundance of Orthoptera species and community descriptors across crop phenological stages and spatial zones in subtropical paddy fields of Northeastern ArgentinaTable 1 long description.

Figure 3

Figure 3. Mean richness (log10) of Orthoptera in subtropical paddy fields of Northeastern Argentina in relation to vegetative (Til = tillering; SE = Stem elongation) and reproductive phenological stages (Flo = flowering; Rip = ripening).

Figure 4

Figure 4. (A) Mean abundance (log10) of the Orthoptera assemblage observed in subtropical paddy fields at Northeastern Argentina in relation to vegetative (Til = tillering; SE = stem elongation) and reproductive phenological stages (Flo = flowering; Rip = ripening). (B) Mean abundance (log10) of adult and nymphal Orthoptera comparing vegetative (Veg) and reproductive (Rep) phenology of rice crops in Northeastern Argentina. Significant differences are indicated by different letters above the bars (post hoc test).Figure 4 long description.

Figure 5

Figure 5. Ranking-abundance curves in two areas of the plots (edge and centre) of paddy fields in Northeastern Argentina. References: Clong (Conocephalus longipes); Avit (Aleuas vitticolis); Bcoc (Belosacris coccineipes); Dexi (Dichroplus exilis); Madsp (Metaleptea adspersa); Etrun (Eucaulopsis truncata); Larg (Leptysma argentina); Tgrac (Tucayaca gracilis), Cang (Coryacris angustipennis); Cysant (Cylindrotettix santarosae); Pacum (Paulinia acuminata); Sxant (Stenacris xanthochlora); Tetri sp. 1 (Tetrigidae sp.); and Aorn (Adimantus ornatissimus).Figure 5 long description.

Figure 6

Figure 6. Venn diagram with the Orthoptera species exclusive to the edges and the centre of the paddy fields, as well as those occurring in both zones. C represents the species turnover value between the two field zones.

Figure 7

Figure 7. Non-metric multidimensional scaling (NMDS) analysis of the orthopteran assemblage in subtropical paddy fields of Northeastern Argentina. Circles represent the replicate sites located in the field-edge areas, whereas squares represent the replicates from the field-centre areas.

Figure 8

Table 2. Trophic level and feeding habits of Orthoptera based on the available literature in subtropical paddy fields of Northeastern Argentina and role as potential pesta of rice, potential natural enemies of pest insectsb, and bioindicatorscTable 2 long description.