Montero-Cedeño, Silvia Lorena ¹ ; Cañarte-Bermúdez, Ernesto Gonzalo ² and Sarmento, Renato Almeida ³

¹✉ Escuela Superior Politécnica Agropecuaria de Manabí, Manuel Félix López, Calceta, Manabí, Ecuador.
²Instituto Nacional de Investigaciones Agropecuarias (INIAP), PO Box 100, Portoviejo, Manabí, Ecuador.
³Universide de Federal de Tocantins (UFT), PO Box 66, Gurupi, TO, Brasil.

2026 - Volume: 66 Issue: 3 pages: 855-867

Original research

Keywords

Abstract

Introduction

Generalist phytoseiid mites are of great importance in the control of several soft-bodied arthropod pests (Döker et al. 2024). However, their reproductive potential is greater when they are fed with pollen. In most cases, prey mites by themselves are insufficient for providing their nutritional requirements, unless a complementary food is present (Broufas & Koveos 2000). Agricultural systems that favour monoculture simplify ecosystems and make it difficult to balance the natural predator-prey relationships (Bermúdez et al. 2010), while diversified systems favour an increase in the density of natural predators by offering microhabitats and alternative food sources (Landis et al. 2000; Nicholls & Altieri 2004). Typhlodromus (Anthoseius) transvaalensis (Nesbitt 1951) (Acari: Phytoseiidae) is a generalist predatory mite, widely distributed in many tropical and subtropical areas of the world, collected from physic nut plants Jatropha curcas L. (Euphorbiaceae), infested with the broad mite Polyphagotarsonemus latus Banks 1904 (Acari: Tarsonemidae) and red mite Tetranychus bastosi Tuttle Baker and Sales 1977 (Acari: Tetranychidae) (Sarmento et al. 2011; Cruz et al. 2013; Pedro-Neto et al. 2013). Little is known concerning food sources for T. (A.) transvaalensis (Horn et al. 2018; Valbuza et al. 2020). Surveys performed on physic nut plants in the State of Tocantins-Brazil revealed a large variety of phytoseiid species associated with phytophagous mites in this crop, with Euseius concordis (Chant), Iphiseiodes zuluagai Denmark & Muma, Typhlodromalus aripo De Leon T. (A.) transvaalensis and Typhlodromalus clavicus Denmark and Muma, standing out for their abundance (Sarmento et al. 2011; Cruz et al. 2013; Saraiva et al. 2014; Marques et al. 2015).

The first record of T. (A.) transvaalensis associated with P. latus corresponds to Cañarte et al. (2017) in spontaneous plants infested with this pest. Momen and Hussein (1999) studied their performance after being fed with different types of prey and castor bean (Ricinus communis L.) pollen. Based on their lifestyle and eating habits (McMurtry & Croft 1997; Luh & Croft 2001), this predator corresponds to a type III generalist, which feeds on prey and pollen, also having thelytokous parthenogenesis reproduction (Ehara & Kishimoto 2007; Kumar,2023). The diversity of feeding habits allows the survival and reproduction of generalist predators, feeding on pollen from several plants (Van Rijn & Tanigoshi,1999; Broufas & Koveos,2000; Vantornhout et al.,2005; Papadopoulos & Papadoulis,2008), exudate of plants, nectar (McMurtry & Croft 1997), fungi (Zemek & Prenerová 1997) and even other types of prey (Nicholls & Altieri 2004). The addition of alternative food to improve the reduction of a pest species through apparent competition has been used in biological control with successful results (Hanna et al. 1997; Van Rijn et al. 2002; Liu et al. 2006).

Several types of pollen have been reported as complementary food in the diet of generalist phytoseiid mites (McMurtry & Croft 1997; Van Rijn & Tanigoshi 1999; Broufas & Koveos 2000; Villanueva & Childers 2004; Bermúdez et al. 2010; Kolokytha et al. 2011; Rodríguez-Cruz et al. 2013). The nutritional value of pollen varies according to the species and botanical family, which influence its biological and reproductive parameters (Van Rijn & Tanigoshi 1999; Emmert et al. 2008; Momen & Abdel-Khalek 2008). James (1993) and Bermúdez et al. (2010) have documented the positive effect of various types of pollen on the biology of phytoseiid mites, particularly in Typhlodromus (Typhlodromus) exhilaratus Ragusa, Typhlodromus (A.) doreenae Schicha, Typhlodromus (T.) pyri Scheuten and Euseius victoriensis (Womersley). In these species, pollen ingestion promoted higher survival and reproductive capacity compared to those fed exclusively on prey such as Panonychus citri (McGregor), Tetranychus urticae Koch and Brevipalpus chilensis Baker. However, the decrease in the predation rate of T. pyri and Euseius hibisci (Chant) in the presence of certain types of pollen, such as Typha latifolia L. (Wei & Walde 1997) is also reported.

A relevant aspect is that the morphology of pollen and the presence of toxic compounds in certain plant species can limit their exploitation by predators (Bermúdez et al. 2010). In this context, mobility and foraging capacity (Slone & Croft 2001), together with the ability of phytoseiid mites to feed on alternative sources, constitute desirable characteristics in a biological control agent. These qualities not only ensure their persistence in the field under conditions of scarcity or absence of prey but also facilitate their mass reproduction in biological control programs (Nomikou et al. 2001; Van Rijn et al. 2002; Gnanvossou et al. 2005).

The predator-prey relationship and its alternative and/or complementary food must be studied for each species of phytoseiid mite. The objective of this research was to evaluate the biological response of the predatory mite T. (A.) transvaalensis after using pollen from Z. mays, J. curcas and R. communis as an alternative and/or complementary food, as well as the effect of three application frequencies of R. communis castor bean pollen. We evaluated the development time of embryonic, post-embryonic, survival, and reproductive parameters in laboratory conditions.

Material and methods

We conducted two experiments in an environmental test room in the Ecology Laboratory at the Federal University of Tocantins (UFT), Gurupi campus. Gurupi-To-Brasil, at 26 ± 1 °C, 70 ± 5% RH and a 12-hour photoperiod. The first experiment aimed to evaluate the biological response of the predatory mite T. transvaalensis after using pollen of Z. mays, J. curcas and R. communis, while the second aimed to evaluate the effect of P. latus and three application frequencies of R. communis pollen on the development time of embryonic, post-embryonic, survival and reproductive parameters.

Typhlodromus transvaalensis rearing

The specimens for this research were obtained from the colony maintained in a breeding room under controlled conditions in the laboratory of the Federal University of Tocantins-Brazil. A colony of T. transvaalensis was maintained in a climate-controlled room (26 ± 1 °C, 70 ± 5% RH, and a 12-hour photoperiod), according to the methodology of Momen (2012) and Pedro-Neto et al. (2013). The specimens were mounted on microscope slides in Hoyer's solution and sent to a specialist at the Federal Rural University of Pernambuco (Brazil) for identification. The colonies were kept on plastic discs (Ø = 9 cm) floating in distilled water inside plastic trays (Cañarte et al. 2017) and fed daily with R. communis pollen (Reis et al. 2007).

Pollen sources

The collection of Z. mays, J. curcas and R. communis pollen was carried out in the morning hours, at the Gurupi campus of the Federal University of Tocantins, Brazil (11º43′45» S, 49º04′07» W and an altitude of 278 m asl) and (26 °C, 72.03% RH and 1184.20 mm of precipitation) (Agrometeorological Station of the Federal University of Tocantins, Gurupi campus). Z. mays pollen was collected from inflorescences, which were placed in paper bags and shaken to obtain pollen. The material was then sieved to eliminate all other plant contamination or arthropods. We selected freshly opened inflorescences to obtain J. curcas pollen; the stamens were rubbed with a short-and-hard bristle brush inside a small black container. Considering that the pollen arrives from the field with humidity between 20 and 30% (Cañarte et al. 2017) the J. curcas pollen was dehydrated in an oven at 50 °C for 48 hours, until the humidity reached the level of approximately 4%, maintaining its properties for up to one year in refrigeration at 4 °C (Bermúdez et al. 2010; Rodríguez-Cruz et al. 2013). The R. communis pollen was obtained from inflorescences cut with a peduncle of 20 to 30 cm in length, immersed in distilled water, and contained in a beaker, which in turn was placed on a white piece of cardboard, where the pollen was deposited. Three days later, the pollen was collected and sieved using a piece of organza fabric, and finally stored in small vials and refrigerated at 10 °C for a maximum period of 30 days (Daud & Feres 2004). To ensure a consistent and reproducible supply during the trials, the three pollen collection and storage methods (fresh, dehydrated, and refrigerated) were employed, which maintained pollen viability, nutritional value, and palatability, while reducing the risk of microbial degradation.

Biological response of T. transvaalensis fed with three types of pollen

The nutritional value of corn (Zea mays), physic nut (Jatropha curcas), and castor bean (Ricinus communis) pollen was evaluated, comparing their effects on the biological, reproductive, and population parameters of the predatory mite T. transvaalensis. Each pollen type was considered an independent treatment, with 30 replicates using plastic discs as experimental units.

Embryonic and post-embryonic development of T. transvaalensis

For the observation of the embryonic period, initially, each of the 30 female T. transvaalensis individuals, of known age, was independently transferred to the experimental units, which consisted of flexible plastic arenas (Ø = 3 cm), arranged in a group of five arenas floating in distilled water, in Petri dishes (15 cm in diameter x 2 cm high) without cover. Water also served as a physical barrier, preventing mites from escaping (Franco et al. 2007; Reis et al. 2007). On the surface of each arena, we placed a small sheet of transparent plastic containing hydrophilic cotton fibre, which served as a refuge and oviposition place for T. transvaalensis, facilitating the observation of the development of the predator (Broufas & Koveos 2000; Rodríguez & Ramos 2004). Each female was observed two times per day until the first egg was laid. Immediately, the females were returned to the rearing units. The eggs were checked twice a day under a stereomicroscope (at 08:00 h and 16:00 h) until the hatch was observed (Rodríguez-Cruz et al. 2013). We recorded the time in days required for the larvae to emerge.

For the evaluation of the post-embryonic development of T. transvaalensis, larvae were maintained in arenas of the same type as previously described. The duration (in days) and survival rates were recorded for the larva, protonymph, and deutonymph stages, as well as for the overall egg-to-adult period. Observations were conducted twice daily, as described above (Rodríguez-Cruz et al. 2013). Molting was confirmed by the presence of exuviae in the arenas (Broufas & Koveos 2000; Rodríguez & Ramos 2004; Bellini et al. 2010).

Once individuals reached the adult stage, observations were conducted once daily at a fixed time (Momen, 2012), recording the duration of the pre-oviposition, oviposition, and post-oviposition periods, as well as adult longevity (Reis et al. 2007). Food was supplied ad libitum according to each treatment (Zea mays, Jatropha curcas, or Ricinus communis pollen) and renewed daily from larval emergence until the death of the last individual. Old pollen was carefully removed to prevent fungal contamination and ensure the continuous availability of high-quality food. In addition, arenas and distilled water in each Petri dish were replaced every 4–5 days (Broufas & Koveos 2000; Momen 2012).

Fertility life table

The adult T. transvaalensis from the post-embryonic development study were maintained in plastic arenas (Ø = 3 cm), grouped in series of five, floating in distilled water inside 15-cm-diameter Petri dishes (Reis et al., 2007). Eggs laid by females were counted and removed every 24 h (Broufas & Koveos 2000; Kolokytha et al. 2011). Food was provided according to the evaluated diet (pollen).

Data for constructing the life table were obtained from the surviving females: 30 for the diet with Z. mays pollen, 23 for J. curcas pollen, and 30 for R. communis pollen, with each female being considered an independent replicate. Reproductive parameters including pre-oviposition, oviposition, and post-oviposition periods were determined, as well as the oviposition rate (total number of eggs laid divided by the duration of the oviposition period) and survival (Kolokytha et al. 2011; Rodríguez-Cruz et al. 2013). The sex ratio was 1:0 (100% females).

The population life-table parameters were calculated using the classical fertility life-table approach described by Andrewartha and Birch (1954), as cited by Silveira Neto et al. (1976), and following the demographic parameters reported by Tanigoshi et al. (1975) and Marafeli et al. (2014), as follows:

R_o = Net reproductive rate (female progeny / female / generation)

R_o = ∑ m_x.l_x

T = Mean generation time (days)

T = ∑ m_x.l_x.x / ∑ m_x.l_x

r_m = Innate capacity for increase in numbers (females / female / day)

r_m = ln(R₀) / T

λ = Finite rate of increase (progeny/female)

λ = e^rm

TD = Time needed for the population to double the number of individuals (days)

TD = ln (2) / r_m

The variation in the estimated life-table parameters was assessed using the jackknife technique described by Maia et al. (2000).

Although iterative solutions of the Euler–Lotka equation are currently commonly used to estimate the intrinsic rate of increase, the classical fertility life-table approach was retained to maintain consistency with the methodology applied throughout the study.

Biological response of T. transvaalensis to a diet of P. latus and three application frequencies of castor bean (R. communis) pollen

In this second experiment, the biological, reproductive, and population parameters of T. transvaalensis- were evaluated, fed with P. latus and R. communis pollen at three frequencies (every 24, 48, and 72 h). Thirty replicates were conducted using plastic discs as experimental units, with a diet consisting of 20 P. latus larvae and fresh R. communis pollen as a supplement. Plastic covers were used to maintain a uniform food supply and to prevent contamination or desiccation (Broufas and Koveos 2000; Kolokytha et al. 2011).

Embryonic, post-embryonic development and survival rate of T. transvaalensis

Thirty T. transvaalensis females of known age were transferred to experimental units consisting of flexible plastic arenas (Ø = 3 cm), grouped in series of five, and floating in distilled water inside 15-cm-diameter Petri dishes, following the methodology described by Broufas and Koveos (2000), Rodríguez and Ramos (2004), and Franco et al. (2007). Each female was observed twice daily until the first egg was laid, after which she was returned to the rearing unit. The eggs were examined twice daily under a stereomicroscope until hatching (Rodríguez-Cruz et al. 2013), and the duration of embryonic development was recorded.

For post-embryonic development, the larvae were maintained in the same type of arena described above. The duration and survival rate of the larval, protonymph, and deutonymph stages, as well as the egg-to-adult cycle, were recorded through twice-daily observations (Rodríguez-Cruz et al. 2013). This methodology was consistent with that reported by Broufas and Koveos (2000), Rodríguez and Ramos (2004), and Bellini et al. (2010).

Once the adult stage was reached, observations were made once a day at a fixed schedule (Momen 2012), recording the pre-oviposition, oviposition, post-oviposition, and longevity of everyone (Franco et al. 2007). The supply of R. communis pollen was adjusted according to each treatment (24, 48, or 72 h), from larval emergence until the death of the last individual. Additionally, 20 P. latus larvae were offered daily at all pollen frequencies. Special attention was paid to removing old pollen to avoid contamination, following the methodology proposed by Broufas and Koveos (2000) and Momen (2012).

Fertility life table under different pollen frequencies and P. latus.

For the pollen frequency experiment, T. transvaalensis adults from the post-embryonic development study were kept in plastic arenas (Ø = 3 cm) (Reis et al. 2007), where predator eggs were counted and removed every 24 h (Broufas and Koveos 2000; Kolokytha et al. 2011). Data for constructing the fertility life table were obtained from the surviving females, totaling 30 per pollen frequency, with each considered an independent replicate. Reproductive parameters such as pre-oviposition, oviposition, and post-oviposition periods, as well as the oviposition rate (total number of eggs laid divided by the oviposition period) and survival rate were determined (Kolokytha et al. 2011; Rodríguez-Cruz et al. 2013). The sex ratio was 1:0 (100% females). Population parameters (Ro, T, rm, λ, and TD) were calculated following the equations described in the previous experiment.

Data analysis

An analysis of variance (ANOVA) was performed on the biological and reproductive parameters of T. transvaalensis in both experiments, followed by Tukey's test (α = 0.05). For females fed different diets and frequencies, the Kaplan–Meier survival curve were fitted, and the results were compared using the Log-rank test (Hosmer et al. 2008). All statistical analyses were performed using R software (R Core Team 2023).

The variation in the fertility life-table parameters was estimated using the jackknife technique described by Maia et al. (2000).

Results

Biological response of T. transvaalensis to three types of pollen

Embryonic, post-embryonic development, and survival rate of T. transvaalensis**. The duration of the egg-to-adult stage differed significantly among the diets (F_2,87 = 75.69, P < 0.01). Development was significantly shorter when the predator was fed on R. communis pollen compared to J. curcas pollen, with the latter resulting in a developmental period more than twice as long (Table 1). The analysis revealed a significant difference among the types of pollen for the incubation period (F_2,87 = 13.78, P < 0.01), with R. communis pollen yielding a lower value than the other two diets (Table 1). In contrast, no significant difference was observed among the pollen types under study during the larval stage (F_2,87 = 0.31, P > 0.05). However, there was a significant effect on both the protonymph (F_2,87 = 64.56, P < 0.01) and deutonymph (F_2,87 = 45.70, P < 0.01) stages.The protonymph stage was shortest when T. transvaalensis was fed with R. communis pollen, differing significantly from J. curcas and Z. mays pollen. Both R. communis and Z. mays pollen significantly reduced the duration of the deutonymph stage relative to J. curcas pollen (Table 1). Notably, all juvenile predators reached adulthood in all cases.

Significant differences were observed in all the reproductive parameters of T. transvaalensis fed on the three types of pollen, including pre-oviposition (F_2,80 = 97.31, P < 0.01), oviposition (F_2,80 = 40.17, P < 0.01), post-oviposition (F_2,80 = 5.53, P < 0.01), oviposition rate (F_2,80 = 25.97, P < 0.01), total eggs per female (F_2,80 = 69.66, P < 0.01), and longevity (F_2,80 = 23.45, P < 0.01). Dietary supplementation with R. communis pollen resulted in the shortest pre-oviposition period and the highest oviposition rate, which differed statistically from Z. mays and J. curcas pollen. Conversely, Z. mays pollen led to the longest oviposition and post-oviposition periods, differing significantly from the other diets. Although the total number of eggs per female did not differ significantly between R. communis and Z. mays pollen, both remained significantly higher than J. curcas pollen, which yielded the lowest mean value. T. transvaalensis exhibited greater longevity on the Z. mays pollen diet compared to the R. communis and J. curcas pollen treatments (Table 1). Overall, the lowest values across all reproductive parameters were consistently recorded for T. transvaalensis fed with J. curcas pollen.

The survival of adult females of T. transvaalensis was influenced by the type of pollen (X² = 28.90, df = 2, P < 0.01). Females fed with Z. mays pollen exhibited a significantly higher survival rate than those provided with R. communis (X² = 12.70, df = 1, P < 0.01) or J. curcas (X² = 41.50, df = 1, P < 0.01) pollen. No significant differences were observed in the survival of females fed with R. communis or J. curcas pollen (X² = 0.90, df .= 1, P = 0.33).

Fertility life table

The analysis of the population parameters of T. transvaalensis revealed significant differences among the three types of pollen across all parameters, including the net reproductive rate (R₀; F_2,118 = 18011, P < 0.0001), mean generation time (T; F_2,118 = 9856, P < 0.0001), innate capacity for increase (r_m; F_2,118 = 20561, P < 0.0001), finite rate of increase (λ; F_2,118 = 20057, P < 0.0001), and doubling time (TD; F_2,118 = 133.1, P < 0.0001) (Table 2).When T. transvaalensis was fed with R. communis pollen, it exhibited superior population parameters, differing significantly from those observed with Z. mays and J. curcas pollen (Table 2). The predator performed best on this diet, demonstrating a higher innate capacity for increase (r_m), a greater net reproductive rate (R₀), and a shorter mean doubling time (TD). The population parameters on Z. mays pollen were similar to those of R. communis. However, with J. curcas pollen, the predator recorded the lowest values for r_m and R₀, along with the longest mean doubling time (TD) (Table 2).

Biological response of T. transvaalensis to a diet of P. latus and three application frequencies of castor bean (R. communis) pollen

Embryonic, post-embryonic development and survival of T. transvaalensis**: No significant differences were observed in the embryonic and post-embryonic stages of T. transvaalensis as a result of the three pollen application frequencies, including the egg stage (F_2,87 = 0.56, P > 0.05), larva (F_2,87 = 1.69, P > 0.05), protonymph (F_2,87 = 2.64, P > 0.05), and deutonymph (F_2,87 = 0.38, P > 0.05) stages. Similarly, the egg-to-adult phase did not differ significantly (F_2,87 = 0.23, P > 0.05). Notably, all juveniles reached adulthood across all three application frequencies of R. communis pollen (Table 3).

The reproductive parameters of T. transvaalensis obtained under the three feeding frequencies revealed significant differences in the pre-oviposition (F_2,87 = 12.43, P < 0.01) and oviposition (F_2,87 = 3.47, P < 0.05) periods. The predator exhibited optimal performance when fed a P. latus larvae diet supplemented with R. communis pollen every 24 and 48 h (Table 3). The duration of the post-oviposition stage also varied significantly among feeding frequencies (F_2,87 = 3.79, P < 0.05), reaching its highest values when the predator was fed every 48 and 72 h. Conversely, the oviposition rate (F_2,87 = 1.27, P > 0.05) and longevity (F_2,87 = 0.54, P > 0.05) of T. transvaalensis remained statistically similar across all treatments. However, the total number of eggs per female differed significantly (F_2,87 = 10.82, P < 0.01), with the highest fecundity observed in the 24-h frequency treatment (Table 3).

The application frequency of R. communis pollen did not significantly influence the survival of adult T. transvaalensis females (X² = 0.90, df = 2, P = 0.65), reflecting highly consistent survival rates among the treatment groups (Fig. 1).

Fertility life table under different pollen frequencies and P. latus

The analysis of the population parameters of T. transvaalensis revealed significant differences among the R. communis pollen feeding frequencies across all parameters, including the net reproductive rate (F_2,130 = 5106, P < 0.0001), T = mean generation time (F_2,130 = 89.36, P < 0.0001), r_m = innate capacity for increase in numbers (F_2.130 = 1929, P < 0.0001), λ = finite rate of increase (F_2,130 = 1948, P < 0.0001) and TD = and doubling time (F_2,130 = 1811, P < 0.0001) (Table 4). Providing R. communis pollen every 24 h to T. transvaalensis fed a P. latus larvae diet resulted in superior population parameters compared to the 48-h and 72-h frequencies. The 24-h frequency was characterized by the highest innate capacity for increase (r_m), the greatest net reproductive rate (R_o), and the shortest mean doubling time (TD). In contrast, the 72-h frequency yielded the lowest values for r_m and R₀, while exhibiting the longest mean doubling time (TD) (Table 4).

Discussion

Typhlodromus transvaalensis developed and reproduced on Z. mays, J. curcas and R. communis pollen. Z. mays and R. communis pollen had the highest nutritional value for T. transvaalensis, a fact reflected by the high activity levels observed during the nymph and adult stages. In contrast, with J. curcas pollen, exhibited limited mobility and clear signs of toxicity. In this regard, James (1993) and Bermúdez et al. (2010) previously described the positive effect of several types of pollen on the development and reproduction of phytoseiid mites closely related to T. transvaalensis. Our findings confirm that the nutritional value of pollen varies significantly across botanical species, directly impacting the biological, reproductive, and survival parameters of predatory mites (Van Rijn & Tanigoshi 1999; Broufas & Koveos 2000; Emmert et al. 2008; Momen & Abdel-Khalek 2008; Kolokytha et al. 2011).

In the present study, T. transvaalensis exhibited favorable biological and reproductive parameters when fed R. communis pollen. Similar results were reported for the phytoseiid Amblyseius herbicolus when provided with the same food source (Rodríguez-Cruz et al. 2013). These findings suggest that R. communis pollen constitutes a suitable nutritional source for different species of predatory mites, likely due to its protein content and other essential nutrients that promote growth and reproduction (Rodríguez-Cruz et al. 2013; Cañarte et al. 2017). The availability of this alternative food resource is critical for maintaining beneficial mite populations during periods of low prey density, which has important implications for biological control programs and the long-term persistence of these predators in agricultural systems (McMurtry & Croft 1997; Gerson et al. 2003).

Zea mays pollen extended the biological cycle of T. transvaalensis while supporting desirable reproductive parameters, such as long periods of oviposition, post-oviposition, longevity, and survival, thereby favoring its permanence in the field. Our results are comparable to those observed for T. athenas (Kolokytha et al. 2011), a successful species related to T. transvaalensis, which exhibited a fertility range like our findings when fed R. communis and Z. mays pollen.

This stands in contrast to J. curcas pollen, which resulted in detrimental biological, reproductive, and population parameters. Although the dehydration of J. curcas pollen in an oven was intended to volatilize toxins, it was insufficient to allow the efficient performance of T. transvaalensis. Several studies attribute this toxicity to the presence of phorbol esters (Wink et al. 1997), which exert a wide range of toxic effects on both vertebrates and invertebrates (King et al. 2009). Previous research has also demonstrated a decrease in the rate of phytoseiid predation in the presence of certain toxic pollens (Wei & Walde 1997). Consequently, plant species with high toxic content do not serve as viable food alternatives (Bermúdez et al. 2010).

Pollen shape and size significantly influence the collection, breakdown, and uptake of food content by Phytoseiidae (Kolokytha et al. 2011). Dietary selection is therefore governed by morphological, physiological, and behavioral factors (Van Rijn & Tanigoshi 1999). These factors may have contributed to the observed differences in the biological performance of T. transvaalensis when fed Z. mays and J. curcas pollen. The extreme size of these grains, in relation to the size of the predator, may have hindered handling and access to the nutritional content. Particularly with J. curcas, the physical difficulty of feeding demanded greater effort from T. transvaalensis, drastically affecting its biological, reproductive, and population parameters, a phenomenon previously noted in studies involving Crotalaria sp. pollen (Rodríguez-Cruz et al. 2013). Conversely, the small size of R. communis pollen facilitated handling and consumption, enabling the predator to reach satisfactory developmental parameters.

The use of pollen as an alternative food for natural enemies has been successfully implemented in biological control programs (Van Rijn et al. 2002). However, there is limited research on the response of predatory mites to pollen application frequency (Broufas & Koveos 2000; van Maanen et al. 2010). The proportional relationship between the reproductive parameters of T. transvaalensis and the application frequency of R. communis pollen suggests efficient utilization of its nutritional richness, which is known to be high in protein (Venzon et al. 2006). Similar successful results were reported by Furtado & Moraes (1998) and Rodríguez-Cruz et al. (2013). Interestingly, application frequency did not influence the oviposition rate of T. transvaalensis. This may be an intrinsic characteristic of the predator when pollen serves as an alternative or complementary food required in low quantities, meaning frequent encounters with pollen are not strictly necessary for reproductive success. According to McMurtry & Croft (1997), Ragusa & Tsolakis (2000), and Luh & Croft (2001), the efficiency of Phytoseiidae lies in their low nutritional requirements, rapid development, high foraging ability, and persistence within plants with low prey infestation. The longevity of T. transvaalensis confirmed the efficient use of pollen at any application frequency, reaching values like other high longevity phytoseiids (Broufas & Koveos 2000; Vargas et al. 2005).

Under circumstances where the predator utilizes alternative food at longer intervals, mobility and searching ability (Slone & Croft 2001), as well as the ability to feed on various sources, are desirable traits for a biological control agent. This ensures environmental persistence when prey is scarce or absent, while also facilitating mass rearing for inclusion in biological control programs (Nomikou et al. 2001; Van Rijn et al. 2002; Bouras & Papadoulis 2005; Gnanvossou et al. 2005).

Finally, the positive influence of R. communis and Z. mays pollen on the reproductive and population parameters of T. transvaalensis provides further support for the potential of agroecological production systems, where the association with crops such as Z. mays and R. communis, among other pollen-supplying plants, could contribute to increasing the diversity and abundance of beneficial species (Landis et al. 2000; Van Rijn et al. 2002; Nomikou et al. 2010). Although generalist phytoseiid mites such as T. transvaalensis prefer prey mites, pollen can provide an important alternative or complementary food source when prey availability is low. Therefore, R. communis may represent a potential supplementary pollen source for supporting predatory mite populations in agricultural systems (Ramakers & Voet 1995, 1996; Van Rijn & Tanigoshi 1999).

However, the use of R. communis as a supplementary pollen source under field conditions requires further evaluation. Future studies should assess whether establishing R. communis as a border or companion plant can provide a sustained pollen resource for T. transvaalensis, while also determining the appropriate plant density and spatial arrangement. These studies should consider predator population dynamics, pest suppression, crop yield, and the costs associated with establishing and maintaining the plants. Such information would help determine the agronomic and economic feasibility of incorporating R. communis into agroecological production systems.

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