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Article

Beneficial Soil Fungi Isolated from Tropical Fruit Crop Systems for Enhancing Yield and Growth in Dragon Fruit in Ecuador

by
Yoansy Garcia
1,*,
Danilo Valdez
2,
Daniel Ponce de Leon
3,
Hypatia Urjilez
1,
Jaime Santos-Pinargote
2 and
Daniel Mancero-Castillo
1,*
1
Research Institute, Agrarian University of Ecuador, Guayaquil 090104, Ecuador
2
Agrarian University of Ecuador, Guayaquil 090104, Ecuador
3
Faculty of Agrarian Sciences, Santa Elena Peninsula State University, La Libertad 16459, Ecuador
*
Authors to whom correspondence should be addressed.
Int. J. Plant Biol. 2025, 16(2), 62; https://doi.org/10.3390/ijpb16020062
Submission received: 18 March 2025 / Revised: 19 April 2025 / Accepted: 30 April 2025 / Published: 5 June 2025
(This article belongs to the Section Plant–Microorganisms Interactions)
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Abstract

:
Rhizospheric fungi are emerging as a critical research component in dragon fruit (Hylocereus spp.) production systems. Introducing beneficial non-native fungi is increasingly common due to their positive effects on plant growth, yield, and pathogen suppression. However, this practice may disrupt soil microbial communities, and commercial isolates often show limited adaptation to local conditions. This study aimed to identify native beneficial soil fungi associated with dragon fruit cultivation on the Ecuadorian coast and evaluate their effect on commercial production. Fungal isolates from four dragon fruit plantations were identified using microscopy and genetic sequencing (ITS, EF-1α, and beta-tubulin). The selected fungi were isolates closely related to Talaromyces tumuli, Trichoderma asperellum, and Paecilomyces lagunculariae. All isolates were tested for pathogenicity using detached cladode assays at the laboratory, and non-phytopathogenic monomorphic cultures were further evaluated in the field under a randomized complete block design consisting of T. asperellum, Talaromyces tumuli, a combination of both, and a water control. The combination of T. asperellum and Talaromyces spp. showed a favorable trend in terms of the plants’ vegetative development. However, inoculating Talaromyces tumuli into the commercial plants exhibited a slow response during the first 20 days of the field evaluations. Still, it resulted in a significant increase in the fruit’s diameter and weight, with increases of 88.23% and 67.64%, respectively, compared to those in the control. T. asperellum presented a lower number of fruits per plant, although it showed an increase in fruit diameter and weight. In conclusion, using the native beneficial fungi T. asperellum and T. tumuli contributes positively to the dragon fruit production system.

1. Introduction

Dragon fruit (Selenicereus spp.), also known locally as pitahaya, is an exotic fruit from the cactus family that has gained importance on a global market due to its increasing demand, particularly in Europe and Asia [1]. Although Latin America has emerged as a central production region, with exponential growth in the past decade, standardized best agricultural practices have not yet been established. A key challenge is the limited availability of technical information on crop management, compelling farmers to rely on empirical knowledge to sustain their production [2].
Applying beneficial microorganisms has improved nutrient use efficiency and reduced the dependence on chemical fertilizers [3]. Certain beneficial fungi such as Trichoderma spp. have been proposed as an alternative to improve soil health and control pathogens. Trichoderma spp. stimulate plant growth and provide protection against pathogenic fungi and nematodes by competing for nutrients and directly inhibiting pathogen activity [4]. Alternatively, some studies have shown that introducing microorganisms can induce significant alterations in the soil’s microbial balance [5]. In this context, Trichoderma, a highly competitive fungus, has been linked to the suppression of pathogenic organisms; however, it may also reduce beneficial microorganisms, potentially disrupting the structure of the resident microbial community [6]. Furthermore, microbial invasions can influence the genetic diversity of resident communities through interactions and horizontal gene transfer, potentially driving genetic modifications and further altering the dynamics of the microbial community [7].
Consequently, several countries have established regulations on using non-native microorganisms in agriculture to mitigate their potential impact on native microbial communities and preserve ecosystem stability [8,9]. European Regulation No. 1107/2009 establishes that microbial inoculants must not cause unacceptable environmental effects and must prioritize the protection of human, animal, and ecosystem health over increased agricultural production [7]. In Ecuador, while there are no specific laws regulating microbial inoculation, Agrocalidad’s guidelines (Art. 18, Section g) explicitly allow the use of suitable microorganisms to improve soil conditions or the availability of nutrients in soil and crops [10]. Although less prescriptive than the European model, this framework aligns with the trend toward sustainable production practices on the Ecuadorian coast.
While pathogenic fungi such as Diaporthe and Colletotrichum have been studied in Ecuador [11,12], beneficial rhizospheric fungi for dragon fruit have not been isolated. This scenario has highlighted the need to isolate and identify native microorganisms with significant effects on crop production. Exploring native fungal species is a practical approach, as locally adapted strains are better suited to specific edaphoclimatic conditions and the rhizosphere of various plant species. Additionally, these native fungi can potentially enhance the efficacy of microbial inoculants [13]. Some native strains have been characterized for their ability to optimize nutrient absorption, improve soil structure, and increase plants’ resistance to stress conditions such as salinity or drought [14]. As a result, a trend has emerged in the commercialization of biofertilizers derived from native microorganisms [15].
Dragon fruit is an emerging crop in Ecuador; however, research on its associated microflora, particularly beneficial fungi, remains limited. Most studies have focused primarily on the identification of pathogenic fungi from genera such as Diaporthe, Colletotrichum, Fusarium, and Aspergillus isolated from vegetative tissues [11,12,16]. Currently, the research is in the diagnostic phase, highlighting the need to develop integrated management systems aiming to enhance crop health and sustainability. This need is further emphasized by the limited knowledge of native beneficial fungi and the emergence of new regulatory frameworks being implemented globally. This study aimed to isolate, identify, and evaluate beneficial fungi associated with dragon fruit cultivation on the Ecuadorian coast, specifically from the rhizosphere soil. The results of this study could contribute to a more profitable and sustainable system for dragon fruit cultivation.

2. Materials and Methods

2.1. Description of the Study Site and Sampling

This study was conducted at 10 sites along the Ecuadorian coast, focusing on representative agricultural land uses, including plantations of dragon fruit (Selenicereus spp.), mango, and banana, each with over five years of continuous production. The selected sites were Hacienda La Fernandita (579,517 m, 9,754,071 m), Finca MAXPRI (574,169 m, 9,755,778 m), Finca PitaSpot (574,586 m, 9,755,395 m), and UAE Experimental Station (666,594 m, 9,766,293 m) for dragon fruit; Hacienda San Judas (622,555 m, 9,824,444 m), Hacienda CC Mangos (582,170 m, 9,735,307 m), and Hacienda La Palma (642,310 m, 9,784,637 m) for mango; and Finca San Vicente (659,360 m, 9,751,472 m), Predios UAE Milagro (658,041 m, 9,764,587 m), and Finca La Envidia (683,999 m, 9,738,608 m) for banana. All of the coordinates are expressed in WGS84 UTM zone 17S (Figure 1). The average annual precipitation ranges from 0 to 1000 mm/year, with mean temperatures between 23 and 25 °C.
Soil and root samples were collected between December 2022 and April 2023 during the vegetative growth and flowering cycles of the dragon fruit plantations. Similarly, the mango and banana plants were in a vegetative physiological stage at the time of sampling. Five vigorous and productive plants were selected at each site to ensure representative sampling. Two soil and root samples (with roots up to 3 mm in diameter and non-suberized) were taken from each selected plant at a depth of 20 cm in the zone of more extensive root development. The sampling was performed with soil at the field capacity. The samples were transported to the laboratories of the Agrarian University of Ecuador in dark sterile containers kept at approximately 4 °C.

2.2. Fungal Isolation

Fungal isolation was carried out using a modified version of the protocol described by Covacevich and Consolo [17]. Soil and root samples were homogenized, and a 10 g subsample was transferred into 90 mL of sterile distilled water and agitated for 20–30 min. A 1 mL aliquot was serially diluted in a test tube containing 9 mL of sterile distilled water, repeating the dilution process five consecutive times. A 1 mL aliquot was plated onto potato dextrose agar (PDA) medium + oxytetracycline from the last three dilutions. The plates were incubated at room temperature (25–30 °C) for approximately one week until characteristic fungal colonies reached the sporulation stage. Fungal isolates were obtained by transferring actively growing hyphal tips onto individual PDA plates. Fungal fitness was assessed using the previously reported correlations with the mycelial growth rate [18,19]. This metric has been considered highly reproducible, and fungal isolates were selected based on the mycelial growth rate previously used by Zhang et al. [20] for effective biocontrol agents and rhizosphere competitors. A minimum growth rate of 5 mm/day was established as a threshold to identify isolates with beneficial potential, as rapid mycelial expansion is often associated with strong competitive abilities, nutrient acquisition efficiency, and antagonistic interactions with pathogens [20,21,22].
To produce monosporic cultures, a uniform-diameter mycelial disk from each isolate grown on PDA was transferred into 10 mL of sterile distilled water and vortexed to obtain a homogeneous suspension. Subsequent dilutions were performed in sterile distilled water to reach a final concentration of 1 × 106 conidia mL−1, estimated using a Neubauer hemocytometer. Then, 200 μ L of the suspension was plated onto PDA. After 24 h of incubation at 25 °C, germinated conidia were observed under a stereomicroscope and individually transferred onto new PDA plates using a flame-sterilized needle. The strains obtained were preserved in glycerol and stored at −18 °C.

2.3. Detached Cladode Pathogenicity Testing

A modified version of the detached stem inoculation protocol described by Mancero Castillo et al. [23] was adapted for Selenicereus cladodes to test the pathogenicity of the fungal isolates. Four cladode sections were inoculated for each fungal isolate, with each being 20 cm long at the laminar flow hood. All dragon fruit cladodes were disinfected, and a 1 cm cross-shaped incision was made on the outer flat side using a sterilized blade. The fungal plugs were then inoculated into each incision and covered with parafilm to assess the pathogenic potential. Cladodes in individual containers were maintained in a growth room with a temperature of 30 ± 2 °C and 60 ± 10% relative humidity. The inoculum plugs were removed after 7 days, and the lesions were evaluated to observe pathogenicity symptoms, including tissue discoloration and lesion formation. A completely randomized design (CRD) was used with four treatments, including a water control. Each treatment was applied to four independent replicates (n = 4). The experimental unit was a detached pitahaya cladode. The presence of symptoms (Yes / No) and lesion diameters were recorded after 7 days. The data were analyzed using a binomial distribution and a logit link function. An isolate was classified as phytopathogenic if two or more replicates showed lesions with discoloration.

2.4. Molecular Identification of Fungal Strains

Genomic DNA was extracted from selected fungal cultures using the DNeasy Plant mini kit (Qiagen, Redwood City, CA, USA) according to the manufacturer’s instructions. The DNA’s integrity and quality were evaluated using microvolume spectrophotometry and agarose gel visualization. The DNA was diluted to a concentration of approximately 20 ng µL−1 for Polymerase Chain Reaction (PCR) amplification using internal transcribed spacers (ITSs) [24] and beta-tubulin, and the translation elongation factor (TEF1-alpha) sequence, widely used for the species-level diagnosis of fungi due to its high conservation and informative variability among fungal taxa, was selected for species-specific identification [25]. PCR amplicons were purified using a PCR Cleanup Kit™ (Qiagen) before sequencing. The DNA sequences were cleaned and assembled using the software Mega (v 11.0.13) [26]. The selected sequences were identified via BLASTn searches against the GenBank database using a threshold of ≥99% identity, which was accessed in April 2024.

2.5. Evaluation of Beneficial Fungi in Commercial Dragon Fruit Production

The effects of the inoculated fungi on dragon fruit production were evaluated at Hacienda La Fernandita, where the soil was classified as a Typic Haplocambid. Randomized soil sampling was carried out in the study plot area at a depth of 20 cm; twelve subsamples were taken and mixed to obtain a composite sample, which was sent for an analysis of the physical and chemical soil fertility at the MEGALAB laboratory. The results are presented in Table 1. The cultivation system followed the “Telegraph Wire” method, in which steel wires supported the plants. The high-density planting arrangement consisted of 0.65 m spacing between the plants and 4.0 m spacing between rows, resulting in a total planting density of 7400 plants per hectare. Fertilization was applied through fertigation, 215 N–67 P2O5–410 K2O kg ha−1. Pruning was performed using the traditional methods to prevent excessive branch overlap in the underlying plants.
The beneficial fungal isolates were evaluated using four treatments: T. asperellum, T. tumuli, the combination of (T. asperellum + T. tumuli), and a water control. The experimental unit consisted of 10 plants arranged in a linear row, with five replications per treatment. Each treatment was applied every 10 days at a concentration of 1 × 106 spores mL−1 [27,28], standardized through direct spore counts using a hemocytometer and adjusted to the desired concentration. The viability of the spores was confirmed before each application using the germination test on potato dextrose agar (PDA) medium, considering only batches with viability above 90%. Soil drench applications were performed a total of five times, starting two months before the final evaluation.
The production variables included pod length (cm), assessed using a measuring tape at 10-day intervals over 50 days; the number of cladodes and fruits per plant, recorded manually at the same intervals; fruit diameter (cm), determined using a Vernier caliper with a 0.05 mm resolution; and fruit weight (g) at harvest, measured using a Sartorius balance with a resolution of 0.01 g.
The evaluation of native beneficial fungi in the commercial dragon fruit under field conditions used a complete randomized block design. The data were processed using a one-factor univariate analysis of variance (ANOVA), and mean comparisons were performed using Tukey’s test (p < 0.05). Before the ANOVA, the assumptions of normality, homoscedasticity, and homogeneity of variance were verified using the Shapiro–Wilk and Levene tests. All assumptions for the parametric analysis were met, validating the use of the statistical model.

3. Results

3.1. Beneficial Fungal Isolate Selection

A total of 45 fungal isolates were cultured on potato dextrose agar (PDA) from the soil and root samples collected from the local fruit plantations and forest areas, including the dragon fruit Selenicereus spp. plantations along the Ecuadorian coast. Of these, 42 isolates were discarded due to them having mycelial growth rates below the established 5 mm/day threshold or morphological characteristics as observed under the microscope consistent with those previously reported for soil-borne fungal pathogens. One Trichoderma spp. isolate was identified from Finca Maxpri, and one Talaromyces sp. was isolated from UAE Station. Subsequent pathogenicity tests using the detached pitahaya cladode sections resulted in the exclusion of an additional isolate from La Fernandita, Paecilomyces lagunculariae, due to the development of tissue discoloration and lesion formation. As a result, only two fungal isolates were retained for further evaluation in the commercial plantations as potential beneficial candidates for dragon fruit production (Table 2).

3.2. Molecular Identification and Characterization

The retained isolates were identified using the ITS, β-tubulin, and tef1-α gene regions. T. asperellum was identified with 100% similarity, while the Talaromyces isolate showed 99.61% identity with T. tumuli, though species-level confirmation was limited due to the taxonomic complexity (Table 3). Morphological characterization under the microscope supported the molecular identification. Monosporic cultures were morphologically characterized under the microscope for each candidate fungal isolate (Figure 2) and identified through genetic sequencing. Molecular identification of these native beneficial fungal candidates using the ITS, β-tubulin, and tef1-α regions indicated significant genetic similarity with Talaromyces tumuli, Trichoderma asperellum, and Paecilomyces lagunculariae.
The fungal isolate belonging to the Talaromyces genus could not be confirmed as the species Talaromyces tumuli due to the high genetic variability and taxonomic complexity of this genus. The diversity of Talaromyces species remains insufficiently characterized, with many species yet to be fully described, making accurate identification challenging despite using the genetic sequences of the ITS and beta-tubulin regions [29,30]. Further phylogenetic analyses adding RPB2 (RNA polymerase II’s second largest subunit) and Calmodulin (CaM) sequences are suggested to resolve this taxonomic ambiguity.

3.3. Pathogenicity Testing of the Fungal Isolates

The detached pathogenicity tests on dragon fruit indicated that T. asperellum (A) showed no adverse reactions in the plant, with favorable wound healing observed at the incision site. Similarly, Talaromyces tumuli (B) showed no harmful effects, and the tissue healing was satisfactory. This indicates that T. asperellum and T. tumuli exhibited no detrimental behavior in the detached pathogenicity tests. In contrast, the fungal strain of Paecilomyces lagunculariae induced black and brown rot, with a mean lesion diameter of 2.4 cm surrounding the incision site in the cladode tissue. The affected areas exhibited yellowish to light brown discoloration, indicating tissue degradation. These symptoms indicate the pathogenic potential of this strain, making it unsuitable for use in dragon fruit (Selenicereus spp.) cultivation (Figure 3). The control wounds (D) showed tissue healing without discoloration, confirming that there was no external contamination at the wound site and that aseptic conditions were maintained during the inoculations.

3.4. Evaluation of Beneficial Fungi in the Plants

The initial characteristics of the soil in which the native beneficial fungi trials were conducted to assess their effect on dragon fruit cultivation are described in Table 1.
Figure 4A shows the growth trends in the dragon fruit pods over 50 days. No significant differences (p < 0.05) were observed among the evaluated treatments, indicating that the fungi did not have a positive or negative effect on the physiological development of the plant pods.
In this regard, Figure 4B shows that the T. asperellum + T. tumuli treatment exhibited the highest number of pods per plant at 10 days, significantly different from T. tumuli (p < 0.05), although similar to the rest of the treatments. No significant differences were found between the beneficial fungi treatments and the control in the remaining time intervals. During the 50-day evaluation period, T. asperellum + T. tumuli showed an increase of 14.3%, the control showed an increase of 16.3%, and T. asperellum recorded 22.1%. On the other hand, T. tumuli, despite having the lowest value at 10 days, managed to reach values comparable to those in the other treatments at 50 days, with an increase of 25.7%, representing 9.4% more than the control. This indicates favorable stimulation of the pod production over time.
In Figure 5A, it can be observed that the control, along with T. asperellum and T. tumuli, maintained the highest number of fruits per plant. This trend remained consistent throughout the 50-day evaluation period. However, T. tumuli showed a significant increase over time, positioning itself among the treatments as that with the best performance in terms of fruit production. On the other hand, T. asperellum presented the lowest number of fruits per plant compared to that in the other treatments (p < 0.05).
The control treatment exhibited the smallest fruit diameter compared to the T. tumuli and T. asperellum treatments (p < 0.05), which increased the fruit diameters by 88.23% and 67.64%, respectively, as shown in Figure 5B. These results indicate that these beneficial fungi could influence nutrition and promote bio-stimulation in dragon fruit cultivation. However, the combination of T. tumuli and T. asperellum did not show a significant effect on fruit diameter. Further research is needed to analyze potential antagonistic interactions and competitive dynamics between these beneficial fungi that may hinder complementary action and reduce overall dragon fruit bio-stimulation.
Table 4 shows the effects of the treatments on the fruit weight at 40 days. The highest average fruit weight was obtained with T. tumuli (752.00 g) (p < 0.05), followed by T. asperellum (616.00 g). The combination of T. asperellum and T. tumuli resulted in a weight of 528.40 g, with no significant difference compared to the control (451.00 g). T. tumuli was the most effective treatment, increasing the fruit weight by 66% relative to that of the control. The combination of both fungi did not provide additional benefits, a trend observed in several of the variables evaluated.
The accelerated increase in the number of fruits per plant along with fruit filling—and consequently weight—was 60% higher compared to that in the control group (451 g), a value consistent with other studies [31]. This suggests that T. tumuli possesses a significant biostimulant capacity, enhancing the development and yield of dragon fruit. This finding aligns with reports on the genus Talaromyces as plant-growth-promoters. For instance, under cadmium stress, (T. pinophilus) MR1 enhanced Arabidopsis biomass by up to 107% [32].
The combination of T. tumuli and T. asperellum did not show additional benefits; the results were statistically comparable to those in the control. Each microorganism individually had better effects on the productive variables. Some studies suggest that co-inoculation may not produce the desired effect due to resource competition between microorganisms [33].

4. Discussion

The findings of this study highlight the relevance of isolating, identifying, and evaluating native fungi with biotechnological potential for the sustainable production of dragon fruit (Selenicereus spp.). The presence of T. tumuli and T. asperellum in the soils and roots of local plantations along the Ecuadorian coast suggests the adaptability of these fungi to the regional edaphoclimatic conditions, reinforcing the concept that native microorganisms are often more efficient and safer for agricultural applications [7]. The potential continental occurrence of T. tumuli highlights the need to investigate this genus further, as this fungal species, first described in 2019 by Peterson and Jurjević [34], belongs to the Talaromyces pinophilus complex, recognized for its industrial and agricultural applications. The genus Talaromyces is notable for producing metabolites with antifungal and antibacterial properties [35,36,37]. This genus has been reported in the rhizosphere of dragon fruit in China [38], South Africa, and the United States [39]. Moreover, T. tumuli has been shown to inhibit the development of other fungi in the rhizosphere [40,41]. The production of antimicrobial exudates by T. tumuli [42,43] gives it a competitive advantage in multiple environments. This ability, combined with its rapid and aggressive growth on PDA medium, supports its prevalence and suggests its potential application in biocontrol or biotechnology. Its isolation is particularly significant in Ecuador, where, according to GBIF [39], this may represent its first continental record. The effect of the T. tumuli treatment on the number of fruits, combined with the largest fruit diameters yielded, aligns with the documented role of Talaromyces spp. in enhancing the morphometric fruit parameters across crops. For instance, in tomato under drought stress, T. omanensis significantly increased the fruit width by 4.28% [44]. Our results suggest that the application of T. tumuli has a significant effect starting 20 days after its application, and similar results were observed with other fungi, such as mycorrhizal treatment during symbiosis [45]. Other authors have also reported that species from the genus Talaromyces act as endophytic fungi with beneficial effects on plants [32,40,44], which is consistent with the results obtained here.
Trichoderma asperellum, on the other hand, was first reported in 1999 [46] and has been extensively documented for its ability to control agricultural diseases through the production of antimicrobial metabolites [47,48]. It also enhances nutrient uptake in plants. More than 1200 occurrences of this species have been reported worldwide, with the highest prevalence in Australia, followed by Asia, Africa, America, and Europe [39].
The phytopathogenicity indicated by the detached cladode tests in the Trichoderma and Talaromyces isolates aligns with studies highlighting the potential of both genera in agriculture, where they have been successfully used as biocontrol agents and soil enhancers [32,36,49]. In contrast, the Paecilomyces lagunculariae isolate reaction contrasts with other species from the same genus, such as Paecilomyces lilacinus, which has been widely recognized for its biocontrol capabilities and ability to promote plant growth by releasing essential nutrients into the soil and producing enzymes that enhance plant health [50].
The significant increase in the number of pods due to the Trichoderma asperellum and Talaromyces tumuli applications represents a crucial production effect, similar to findings from other studies in tomato and rice crops [44,51]. These fungi can stimulate plant growth by improving the availability of essential nutrients such as phosphorus and nitrogen, resulting in greater efficiency in their absorption. Furthermore, they increase the production of siderophores, compounds that play a role in iron acquisition, which promotes plant development, particularly under stress conditions [37]. In mixed microbial systems in the rhizosphere, competition for essential nutrients such as iron and phosphorus can significantly influence the performance of beneficial fungi [52]. Both Talaromyces spp. [53] and T. asperellum [54] are known to produce siderophores—iron-chelating compounds that facilitate iron uptake under limiting conditions. However, siderophore-mediated competition may result in inhibitory effects if one species more effectively sequesters the available iron, depriving the other of this critical micronutrient. Similarly, phosphate solubilization via organic acid production may lead to direct competition if both fungi rely on similar pathways or substrates [55]. This competition can reduce the availability of solubilized nutrients for plant uptake or disrupt the ecological balance required for synergistic action. Moreover, pH modulation or the secretion of secondary metabolites during nutrient solubilization may interfere with the metabolic activity or growth of the companion fungus [56]. These potential interactions highlight the importance of evaluating compatibility and resource partitioning when developing multi-agent biocontrol formulations.
Although both T. tumuli and T. asperellum exhibited positive results in productive variables such as fruit weight and diameter, the combined application of these beneficial fungi did not yield synergistic effects. This outcome aligns with the hypothesis of ecological competition among microorganisms in similar niches [33]. Long-term field trials under varying edaphoclimatic conditions are necessary to refine the application strategies and determine when it is more appropriate to apply each strain independently.
Moreover, the identification of the Paecilomyces lagunculariae isolate’s production of pathogenicity symptoms underscores the need to include pathogenicity assays before validating any isolates for agronomic use, even those from genera traditionally considered beneficial [57]. The high genetic variability within the Talaromyces genus also poses the challenge of the need to use additional molecular markers such as RPB2 and calmodulin for precise taxonomic identification [29,30].

5. Conclusions

These results indicate a procedure for isolating and identifying beneficial native fungi for dragon fruit cultivation, such as T. tumuli and T. asperellum. The application of these fungi positively impacts vegetative development and fruit quality, highlighting the importance of employing resident microorganisms to enhance agricultural production without disrupting the soil’s microbiological communities. T. tumuli showed a significant increase in the diameter and weight of the fruit, highlighting its potential to boost productivity. Moreover, the combination of T. asperellum and T. tumuli promoted plant development, but their individual applications obtained better results. This study demonstrates that native fungi are an effective and sustainable tool for optimizing dragon fruit production and reducing the risks of introducing foreign species.

Author Contributions

Conceptualization: Y.G. and D.M.-C. Methodology: D.M.-C., J.S.-P. and Y.G. Software: Y.G. and D.P.d.L. Formal analysis: D.V. and D.M.-C. Investigation: H.U. Resources: D.V. Data curation: Y.G. Writing—original draft preparation: Y.G. and D.M.-C. Writing—review and editing: Y.G. and D.M.-C. Visualization: D.P.d.L. Supervision: Y.G. Project administration: Y.G. Funding acquisition: Y.G. and D.M.-C. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by la Universidad Agraria del Ecuador (Resolution No. 441-2022), Ecuador.

Data Availability Statement

The data presented in this study are available on request from the corresponding author due to institutional and national regulations.

Acknowledgments

The authors thank Gabriel Alfonzo for his collaboration in this research’s development.

Conflicts of Interest

The authors declare that this research was conducted without any commercial or financial relationships that could be construed as potential conflicts of interest.

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Figure 1. A map of the sampling locations along the Ecuadorian coast where soil and root samples were collected for the isolation of beneficial fungi associated with dragon fruit (Selenicereus spp.), mango (Mangifera indica), and banana (Musa spp.) plantations.
Figure 1. A map of the sampling locations along the Ecuadorian coast where soil and root samples were collected for the isolation of beneficial fungi associated with dragon fruit (Selenicereus spp.), mango (Mangifera indica), and banana (Musa spp.) plantations.
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Figure 2. Monosporic cultures of (A) Talaromyces tumuli, (B) Trichoderma asperellum, and (C) Paecilomyces lagunculariae; (D,E) conidiophores, conidia, and septate hyphae of T. tumuli; (F,G) conidiophores, conidia, and septate hyphae of T. asperellum; and (H,I) oogonium and oospores of Paecilomyces lagunculariae.
Figure 2. Monosporic cultures of (A) Talaromyces tumuli, (B) Trichoderma asperellum, and (C) Paecilomyces lagunculariae; (D,E) conidiophores, conidia, and septate hyphae of T. tumuli; (F,G) conidiophores, conidia, and septate hyphae of T. asperellum; and (H,I) oogonium and oospores of Paecilomyces lagunculariae.
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Figure 3. Pathogenicity testing of fungal isolates on dragon fruit. (A) Trichoderma asperellum, (B) Talaromyces tumuli, (C) Paecilomyces lagunculariae, and (D) the control.
Figure 3. Pathogenicity testing of fungal isolates on dragon fruit. (A) Trichoderma asperellum, (B) Talaromyces tumuli, (C) Paecilomyces lagunculariae, and (D) the control.
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Figure 4. The effect of beneficial fungi and their combinations on the cladode length (A) and cladode number per plant (B) in dragon fruit. Different letters above the bars indicate statistically significant differences among the treatments at each evaluation time according to Tukey’s test (p < 0.05). The lines on each bar represent the standard deviation (± SD). n.s. = not significant.
Figure 4. The effect of beneficial fungi and their combinations on the cladode length (A) and cladode number per plant (B) in dragon fruit. Different letters above the bars indicate statistically significant differences among the treatments at each evaluation time according to Tukey’s test (p < 0.05). The lines on each bar represent the standard deviation (± SD). n.s. = not significant.
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Figure 5. The effect of beneficial fungi and their combination on the number of dragon fruits per plant (A) and fruit diameter (B). Different letters above the bars indicate statistically significant differences among the treatments at each evaluation time, as determined by Tukey’s test (p < 0.05). The lines on each bar represent the standard deviation (±SD). n.s. = not significant.
Figure 5. The effect of beneficial fungi and their combination on the number of dragon fruits per plant (A) and fruit diameter (B). Different letters above the bars indicate statistically significant differences among the treatments at each evaluation time, as determined by Tukey’s test (p < 0.05). The lines on each bar represent the standard deviation (±SD). n.s. = not significant.
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Table 1. Initial characteristics of the soil in which the effect of the fungal isolates on dragon fruit cultivation was evaluated.
Table 1. Initial characteristics of the soil in which the effect of the fungal isolates on dragon fruit cultivation was evaluated.
CharacteristicResultCharacteristicResult
pH6.60Sulfur (ppm)11.00
Organic matter (%)1.70Boron (ppm)1.14
CEC (meq 100 g−1)33.30Copper (ppm)7.90
Phosphorus (ppm)11.00Iron (ppm)485.00
Potassium (meq 100 g−1)0.79Manganese (ppm)38.00
Calcium (meq 100 g−1)19.66Molybdenum (ppm)0.03
Magnesium (meq 100 g−1)5.80Zinc (ppm)2.20
Sodium (meq 100 g−1)0.41Microbial biomass (ppm)668.00
Exchangeable Al (meq 100 g−1)<0.01Potentially mineralizable nitrogen (kg N ha−1)19.00
TextureLoamy Clay
Notes: Chemical determinations: B (hot water extractant); P (Olsen extractant, sodium bicarbonate); Ca, K, Mg, and Na (1M ammonium nitrate extractant); Cu, Fe, and Zn (0.05M disodium EDTA extractant); Mn (1M ammonium acetate and quinol extractant); Mo (Tamm’s reagent); soil organic matter (Dumas combustion); pH-H2O (soil/water ratio 1:2.5); S (calcium phosphate tetrahydrogendiorthophosphate extractant); exchangeable Al (calcium sulfate extractant); and microbial biomass (calculated from CO2 (Solvita CO2-Burst)).
Table 2. Selection of beneficial fungal isolates from fruit rhizosphere root and soil samples based on mycelial growth rate and pathogenicity testing on cladode sections.
Table 2. Selection of beneficial fungal isolates from fruit rhizosphere root and soil samples based on mycelial growth rate and pathogenicity testing on cladode sections.
LocationTotal Fungal IsolatesDiscarded Mycelial Growth Rate < 5 mm/dayPhytopathogenic Isolates *Selected Beneficial Isolates
Finca La Fernandita121110
Finca Maxpri9801
Finca PitaSpot8800
UAE Station161501
* Fungal isolates showing lesions on more than two cladode replicates were classified as phytopathogenic and excluded. A minimum mycelial growth rate threshold of 5 mm/day on PDA was used for selection.
Table 3. Molecular identification of fungal isolates from dragon fruit cultivation soil on the Ecuadorian coast.
Table 3. Molecular identification of fungal isolates from dragon fruit cultivation soil on the Ecuadorian coast.
OrganismLocation (WGS84 UTM 17S)% IdentityAccession No. *
Paecilomyces lagunculariae579517, 975407199.55NR145144.1
Talaromyces tumuli666594, 976629399.61NR_165528.1
Trichoderma asperellum574169, 9755778100.00NR_130668.1
* Accession number obtained using the NCBI BLAST tool (https://blast.ncbi.nlm.nih.gov/Blast.cgi, accessed on 29 April 2025).
Table 4. The effect of the native fungi (T. tumuli and T. asperellum) and their combination on dragon fruit weight in a high-density production system on the Ecuadorian coast.
Table 4. The effect of the native fungi (T. tumuli and T. asperellum) and their combination on dragon fruit weight in a high-density production system on the Ecuadorian coast.
TreatmentsFruit Weight at 40 Days (g)
T. tumuli 752.00 ± 41.77  a
T. asperellum 616.00 ± 45.60  b
T. asperellum + T. tumuli 528.40 ± 23.30  c
control 451.00 ± 35.60  c
p-value1.395 × 10−6
Coefficient of variation (%)7.71
Different letters indicate significant differences among treatments according to Tukey’s test (p < 0.05).
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Garcia, Y.; Valdez, D.; Ponce de Leon, D.; Urjilez, H.; Santos-Pinargote, J.; Mancero-Castillo, D. Beneficial Soil Fungi Isolated from Tropical Fruit Crop Systems for Enhancing Yield and Growth in Dragon Fruit in Ecuador. Int. J. Plant Biol. 2025, 16, 62. https://doi.org/10.3390/ijpb16020062

AMA Style

Garcia Y, Valdez D, Ponce de Leon D, Urjilez H, Santos-Pinargote J, Mancero-Castillo D. Beneficial Soil Fungi Isolated from Tropical Fruit Crop Systems for Enhancing Yield and Growth in Dragon Fruit in Ecuador. International Journal of Plant Biology. 2025; 16(2):62. https://doi.org/10.3390/ijpb16020062

Chicago/Turabian Style

Garcia, Yoansy, Danilo Valdez, Daniel Ponce de Leon, Hypatia Urjilez, Jaime Santos-Pinargote, and Daniel Mancero-Castillo. 2025. "Beneficial Soil Fungi Isolated from Tropical Fruit Crop Systems for Enhancing Yield and Growth in Dragon Fruit in Ecuador" International Journal of Plant Biology 16, no. 2: 62. https://doi.org/10.3390/ijpb16020062

APA Style

Garcia, Y., Valdez, D., Ponce de Leon, D., Urjilez, H., Santos-Pinargote, J., & Mancero-Castillo, D. (2025). Beneficial Soil Fungi Isolated from Tropical Fruit Crop Systems for Enhancing Yield and Growth in Dragon Fruit in Ecuador. International Journal of Plant Biology, 16(2), 62. https://doi.org/10.3390/ijpb16020062

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