# Development, reproduction and life table parameters of Tetranychus phaselus Ehara (Acari: Tetranychidae) on two bean host plants

Chen Chen, Wei Pan, Xiaoqian Tong, Jing Song, Yang Pan, Peng Zhou

► To cite this version:

Chen Chen, Wei Pan, Xiaoqian Tong, Jing Song, Yang Pan, et al.. Development, reproduction and life table parameters of Tetranychus phaselus Ehara (Acari: Tetranychidae) on two bean host plants. Acarologia, 2025, 65 (1), pp.197-203. 10.24349/men5-cyrn. hal-05006784

HAL Id: hal-05006784

https://hal.science/hal-05006784v1

Submitted on 26 Mar 2025

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
# Development, reproduction and life table parameters of *Tetranychus phaselus* Ehara (Acari: Tetranychidae) on two bean host plants

Chen Chen<sup>a</sup>, Wei Pan<sup>a</sup>, Xiaoqian Tong<sup>b</sup>, Jing Song<sup>a</sup>, Yang Pan<sup>a</sup>, Peng Zhou<sup>a</sup>

## Original research

### ABSTRACT

Investigations into the life history characteristics of pests on host plants can provide important information on pest forecast and management. *Tetranychus phaselus* Ehara is a polyphagous pest distributed in China, Russia, Japan and Korea. *Glycine max* and *Phaseolus vulgaris* are two most economically important host crops of *T. phaselus*, and the increasing planting area of the two plants may facilitate the occurrence of the pest. Here we investigated the development, survival, reproduction and life table parameters of *T. phaselus* on *G. max* and *P. vulgaris* and how these traits vary between the two plants. We showed that the developmental period from egg to adult was significantly shorter on *G. max* than on *P. vulgaris* for both males and females. *Tetranychus phaselus* had significantly higher egg hatchability and egg-to-adult survival on *G. max*. *Tetranychus phaselus* had different lifetime reproductive pattern on the two host plants and produced significantly more daily and lifetime number of eggs on *G. max* than on *P. vulgaris*. We showed that females fed on *G. max* had significantly higher intrinsic rate of increase (*r*<sub>m</sub>) and net reproductive rate (*Ro*) and shorter generation time (*T*) and doubling time (*Dt*). These results suggest that *G. max* (soybean) is a more favorable host plant for *T. phaselus*, and the increasing planting area of these beans particularly soybeans may facilitate the occurrence of *T. phaselus*.

**Keywords** bean plant; development; fecundity; life table; spider mite

### Introduction

Investigations into how host plants affect life history characteristics of pest herbivores help better understand their abundance and distribution, the knowledge of which is important for pest forecast and integrated pest management programs (Santamaría *et al.* 2018). Host plants vary in chemical traits such as toxins, nutritional quality and quantity, and physical traits such as tissue toughness, which affect development, reproduction, survival and population growth of herbivores (de Lima *et al.* 2017; Golizadeh *et al.* 2017; Santamaría *et al.* 2018; Puspitarini *et al.* 2021; Segura-Martínez *et al.* 2023).

Many spider mites in the family Tetranychidae (Acari: Prostigmata) are highly polyphagous pests worldwide. Studies have shown that different host plants affect development, reproduction and demographic parameters of spider mites such as *Tetranychus urticae* Koch, *T. evansi* Baker & Pritchard (Santamaría *et al.* 2018), *T. ludeni* Zacher (Adango *et al.* 2006), and *T. phaselus* Ehara (Ito *et al.* 2017). *Tetranychus phaselus* is a polyphagous pest distributed in China, Russia, Japan, and Korea, which is currently considered as a minor pest (Ehara, 1960; Zhao

<figure><img src="image_2.png" /></figure>

<footnote><sup>a</sup> School of Life Sciences, Anqing Normal University, Anqing 246133, China.</footnote>

<footnote><sup>b</sup> School of Life Sciences, Anhui University, Hefei 230031, China.</footnote>
et al. 2013; Gotoh et al. 2015; Jin et al. 2018). It has been recorded to appear on at least 16
host plants including *Gossypium* spp. (Malvales: Malvaceae), *Humulus lupulus* L. (Rosales:
Cannabaceae), *Fatoua villosa* (Thunb.) Nakai (Rosales: Moraceae) (Jin et al. 2018; Migeon
and Dorkeld 2024), among which the soybeans *Glycine max* Merr. (Fabales: Fabaceae) and
the common beans *Phaseolus vulgaris* L. (Fabales: Fabaceae) are two most economically
important crops cultivated worldwide. *Glycine max* and *P. vulgaris* belong to Fabaceae, which
are considered favorable host plants of many *Tetranychus* mites (van den Boom et al. 2003;
Ito et al. 2017; Segura-Martínez et al. 2023; Weerawansha et al. 2023; Zhou et al. 2024).
*Glycine max* has more nutritional components such as protein than *P. vulgaris* (Alcázar-Valle
et al. 2020; Li et al. 2024) and may be a more suitable host plant for *T. phaselus*.

The increasing planting area of host plants may alter species composition and facilitate
occurrence of particular pests (Jin et al. 2018). Due to its nutritional and health benefits, there
is an increase of demand and cultivation of Fabaceae beans (FAO 2025). For example, the
planting area of soybean increased from 6.8 million hectares in 2015 to 10 million hectares
in 2022 in mainland China (NBSC 2025), and there is a steady increase of planting area of
common beans in temperate countries including China (Wu et al. 2020), which may facilitate
the occurrence of *T. phaselus*. Numerous studies used common bean plants as host species to
test the performance of spider mites, for example, Gotoh et al. (2015) tested the fitness of four
mites on *P. vulgaris* in response to different temperatures. However, the performances of many
spider mites including *T. phaselus* on soybean plants are not clear. It is also not clear whether *T.
phaselus* performs better on soybean than on common bean. Here we investigated development,
reproduction, survival and life table parameters of *T. phaselus* on *G. max* and *P. vulgaris* and
how these traits vary between the two plants. Knowledge generated could contribute to pest
forecast and integrated pest management programs for controlling this mite in soybean and
common bean growing regions.

**Material and methods**

**Mite colony and experimental conditions**

*Tetranychus phaselus* adults were collected on *Morus alba* L. in Anqing, Anhui province, China, and the colony was established and maintained using 3- to 5-week-old common bean *P. vulgaris* plants grown in pots. *Phaseolus vulgaris* and *G. max* seeds were purchased from local market and were grown in pots. The first expanded leaves cut from 1- to 2-week-old plants were used for experiments. The colony was maintained and the experiments were carried out under the environmental conditions of 24 ± 1 °C, 60 ± 10% RH, and 14:10 h (light:dark) photoperiod in a walk-in climate chamber.

**Immature development**

Forty adult females were randomly selected from the colony and placed on a *P. vulgaris* leaf square (3 × 3 cm). Females were allowed to lay eggs for eight hours and then removed from the leaf square. The eggs were individually transferred onto either a *P. vulgaris* or a *G. max* leaf circle (1 cm in diameter) using a fine brush. Only data of diploid eggs were collected in this procedure. In spider mites, mated mothers produce haploid (giving rise to sons) and diploid eggs (developing to daughters) whereas virgin mothers produce haploid eggs only. Mated females usually produce female-biased sex ratios (e.g., 3:1). To reduce workload, virgin females were used for producing haploid eggs. Fifty female deutonymphs were randomly selected, placed on a *P. vulgaris* leaf square (3 × 3 cm) and allowed to develop into 3-d-old adults. Forty adult virgin females were randomly selected and transferred onto a new leaf square and allowed to lay eggs for eight hours. The adults were then removed and each of the eggs was transferred onto either a *P. vulgaris* or a *G. max* leaf circle (1 cm in diameter) using a fine brush. The individuals were checked every 24 hours, and the developmental period of
eggs, larvae, protonymphs, deutonymphs of males and females were recorded. A total of 60 replicates, i.e., 30 (15 males and 15 females) replicates for each host plant were obtained.

## Reproduction, longevity, and offspring survival

Male and female deutonymphs were randomly selected from the colony and each individual was placed on a leaf circle (1.5 cm in diameter) to develop into 1-d-old adults. To obtain mated females, a female and a male were paired on a new leaf circle for 24 hours and then the male was removed. The mating status of a female was checked by examining whether she produced at least a daughter, and only mated females were used for experiment. A new leaf circle was replaced for the female every 24 hours until the female died. The number of eggs laid on each leaf circle was recorded and the eggs were allowed to develop into adults. The number of eggs hatched, the number of offspring that developed into adults, sex ratio, female oviposition period and female longevity were also recorded. Seventeen and 18 replicates were prepared for *G. max* and *P. vulgaris*, respectively.

## Statistical analysis

All data were analyzed using SAS software (SAS 9.4, SAS Institute Inc., NC, USA). Data distribution was tested using Shapiro-Wilk test (UNIVARIATE Procedure). Data on the male total developmental period (egg to adult) were normally distributed and analyzed using the Student's t-test (TTEST Procedure). Other data were not normally distributed and analyzed using the nonparametric Wilcoxon signed-rank test (NPARIWAY Procedure). A Gaussian functional model according to Archontoulis and Miguez (2015): $y = ae^{(-0.5[\ln(x/x_0)/b]^2)}$ was used to fit the daily number of eggs and daughters for each treatment, where x is the age of females (days), a is the peak at time $x_0$ (female age), b is the coefficient controlling the width of the peak. The model allowed us to compare reproduction patterns between treatments. If the 95% confidence limits of a given parameter overlap, then there is no significant difference between treatments.

The life table parameters including the intrinsic rate of increase, the net reproductive rate, mean generation time, and doubling time were calculated. The life table parameters (Jervis *et al.* 2005) were calculated using the data on daily survival and daughter production of each female. The intrinsic rate of natural increase (r) was calculated by solving the Lotka-Euler equation: $\sum e^{-rx}l_xm_x = 1$, where x is the pivotal age, $l_x$ is the proportion of females surviving to age x, and $m_x$ is the number of daughters produced per female at age x. The net reproductive rate ($R_0 = \sum l_xm_x$, daughters/female/generation), doubling time [$D_t = \log_e(2)/r$, days], and generation time [$T = \log_e(R_0)/r$, days] were also calculated. The bootstrap method with 50,000 bootstrap samples was used to calculate the pseudo-values of a given parameter and the paired-bootstrap test was employed to compare between two treatments (TTEST Procedure).

## Results and discussion

Our results show that *T. phaselus* performed differently on *G. max* and *P. vulgaris* in terms of the development time, reproduction and life-table parameters. Host plants have had a strong influence on the biological parameters of several species of tetranychids (de Lima *et al.* 2017; Golizadeh *et al.* 2017; Santamaría *et al.* 2018; Puspitarini *et al.* 2021; Segura-Martínez *et al.* 2023). The total developmental period of *T. phaselus* from egg to adult was significantly shorter on *G. max* than on *P. vulgaris* for both males and females. In spider mites, the male has a shorter immature stage than the female. Additionally, males typically stop growing after the protonymphal stage and are smaller than females upon emergence as adults (Li and Zhang 2018). These differences may result in divergent nutritional demands across developmental stages between males and females. Our results show that females on *G. max* had significantly shorter developmental period at deutonymph stage, whereas males had significantly shorter
developmental period at the larva and protonymph stage (Table 1). Similarly, different growth rates on different host plant between males and females was also found in the red spider mite *T. merganser* Boudreaux (Segura-Martínez *et al.* 2023). Our results show that *T. phaselus* had significantly higher egg-to-adult survival on *G. max* than on *P. vulgaris* (Table 2). Specifically, egg hatchability was significantly higher on *G. max* than on *P. vulgaris* but larva-to-adult survival was similar between the two host plants indicating that egg stage is more affected by host plants (Table 2). It is widely reported that mothers of spider mites dynamically adjust their resource provisioning to offspring (e.g., egg size) in response to environmental conditions such as host plant quality variations (Dahirel *et al.* 2019), population density fluctuations (Weerawansha *et al.* 2022), and changes in mating status (Zhou *et al.* 2018). The differences in egg hatchability between the two bean plants may be attributed to variations in host plant quality (e.g., leaf nutrient composition) and maternal effects. Faster developmental rate may allow a shorter life cycle and faster population growth rate, and together with higher offspring survival, these features may lead to larger population size on *G. max*.

*Tetranychus phaselus* females had similar oviposition period and longevity on the two host plants but produced significantly more daily and lifetime number of eggs on *G. max* than on *P. vulgaris* (Table 2). The higher reproductive output is also illustrated in Figure 1 where the peaks of daily number of eggs and daughters produced were significantly higher and delayed when fed on *G. max* than those fed on *P. vulgaris* (Figure 1). Ito *et al.* (2017) reported that *T. phaselus* produced about 11 eggs on the favorable host plant *P. vulgaris* within the first five days, which is similar with present study (about 13 eggs within five days). Our results suggest that *G. max* is a more favorable host plant for *T. phaselus*, probably because it is higher in some nutritional components such as protein. Similarly, higher nitrogen content leads to faster

<table><caption>**Table 1.** Developmental period of female and male *T. phaselus* at different stages reared on *G. max* and *P. vulgaris*.</caption><thead><tr><th>Treatment</th><th>Egg</th><th>Larva</th><th>Protonymph</th><th>Deutonymph</th><th>Total (egg to adult)</th></tr></thead><tbody><tr><td>Female</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><i>G. max</i></td><td>4.73 ± 0.25</td><td>2.20 ± 0.20</td><td>2.33 ± 0.19</td><td>3.07 ± 0.15</td><td>12.33 ± 0.36</td></tr><tr><td><i>P. vulgaris</i></td><td>4.67 ± 0.13</td><td>2.67 ± 0.39</td><td>3.13 ± 0.39</td><td>5.47 ± 0.58</td><td>15.93 ± 1.14</td></tr><tr><td>χ<sup>2</sup> (df)</td><td>0.1184 (1)</td><td>0.4642 (1)</td><td>1.7632 (1)</td><td>12.9261 (1)</td><td>9.7195 (1)</td></tr><tr><td>P</td><td>0.7308</td><td>0.4956</td><td>0.1842</td><td>0.0003</td><td>0.0018</td></tr><tr><td>Male</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><i>G. max</i></td><td>5.20 ± 0.14</td><td>1.47 ± 0.13</td><td>2.47 ± 0.26</td><td>3.80 ± 0.45</td><td>12.93 ± 0.46</td></tr><tr><td><i>P. vulgaris</i></td><td>5.40 ± 0.13</td><td>3.73 ± 0.21</td><td>3.53 ± 0.29</td><td>4.27 ± 0.33</td><td>16.93 ± 0.45</td></tr><tr><td>χ<sup>2</sup> (df)</td><td>0.9218 (1)</td><td>21.7107 (1)</td><td>5.8975 (1)</td><td>1.8145 (1)</td><td>t = 6.18 (df=28)</td></tr><tr><td>P</td><td>0.3370</td><td>&lt; 0.0001</td><td>0.0152</td><td>0.1780</td><td>&lt; 0.0001</td></tr></tbody></table>

<table><caption>**Table 2.** Reproduction, offspring survival, and adult longevity of *T. phaselus* reared on *G. max* and *P. vulgaris*.</caption><thead><tr><th>Host plant</th><th><i>G. max</i></th><th><i>P. vulgaris</i></th><th>χ<sup>2</sup> (df)</th><th>P</th></tr></thead><tbody><tr><td>No. eggs</td><td>64.59 ± 8.83</td><td>24.89 ± 3.58</td><td>9.23 (1)</td><td>0.0024</td></tr><tr><td>Oviposition period</td><td>14.59 ± 1.59</td><td>13.72 ± 1.49</td><td>0.11 (1)</td><td>0.7408</td></tr><tr><td>Daily fecundity (No. eggs/female/day)</td><td>3.47 ± 0.36</td><td>1.58 ± 0.22</td><td>11.56 (1)</td><td>0.0007</td></tr><tr><td>Egg hatchability (%)</td><td>98.59 ± 0.41</td><td>93.11 ± 1.83</td><td>4.26 (1)</td><td>0.0390</td></tr><tr><td>Larva-to-adult survival</td><td>93.49 ± 1.60</td><td>88.37 ± 2.93</td><td>1.72 (1)</td><td>0.1896</td></tr><tr><td>Egg-to-adult survival</td><td>92.19 ± 1.69</td><td>82.16 ± 3.10</td><td>6.56 (1)</td><td>0.0104</td></tr><tr><td>Daughters (%)</td><td>79.07 ± 2.82</td><td>70.05 ± 4.72</td><td>2.21 (1)</td><td>0.1373</td></tr><tr><td>Longevity</td><td>17.29 ± 1.39</td><td>16.44 ± 1.45</td><td>0.08 (1)</td><td>0.7785</td></tr></tbody></table>
<figure><img src="image_3.png" /><figcaption>**Figure 1.** The estimated patterns of daily number of eggs (a) and daughters (b) produced by *T. phaselus* on *G. max* and *P. vulgaris*. A Gaussian functional model: $y = ae^{(-0.5|\ln(x/x_0)/b|^2)}$ was used to fit the data, where *x* is the age of females (days), *a* is the peak of curves at time $x_0$ (female age), *b* is the coefficient controlling the width of the peak. The "peak", "width", and "time" in the figures represent *a*, *b*, and $x_0$ of the model, respectively.</figcaption></figure>

development and higher fecundity in *Panonychus ulmi* (Koch) (Breukel and Post 1959), *T. ludeni* (Adango et al. 2006), and *T. urticae* (Wermelinger et al. 1991).

Life table parameters indicate the population growth rates of species in the present generation and subsequent generations and thus a reliable tool to assess the performance of phytophagous arthropods on different host plants. We show that females fed on *G. max* had significantly higher intrinsic rate of increase ($r_m$) and net reproductive rate (*Ro*) and shorter generation time (*T*) and doubling time (*Dt*) (Table 3). The values of life table parameters ($r_m$, *Ro*, *T*, and *Dt*) obtained from the two host plants in our study fell within the ranges documented for *Tetranychus* species in the literature (Kasap 2004; Adango et al. 2006; Sedaratian et al. 2011; Golizadeh et al. 2017; Zhou et al. 2021; de Lima et al. 2022; Ristyadi et al. 2022; Segura-Martínez et al. 2023). The higher $r_m$ on soybean may due to higher reproductive rate (Figure 1) and shorter generation time (Golizadeh et al. 2017; Osman et al. 2019), which is suggested to be the most appropriate index to evaluate the performance of pests on host plants (Golizadeh et al. 2017). These results further suggest that *G. max* is more susceptible to *T. phaselus* attack than *P. vulgaris*. Life table parameters can reflect the occurrence of pests on host plants in the field (Adango et al. 2006; de Lima et al. 2017), and thus provide essential

<table><caption>**Table 3.** The estimated mean (± SE) life table parameters of *T. phaselus* on *G. max* and *P. vulgaris*.</caption><thead><tr><th>Treatment</th><th>r<sub>m</sub> (daughters/female/day)</th><th>Ro (daughters/female/generation)</th><th>T (days)</th><th>Dt (days)</th></tr></thead><tbody><tr><td>G. max</td><td>0.2010 ± 0.0001</td><td>48.41 ± 0.44</td><td>19.30 ± 0.04</td><td>3.45 ± 0.00</td></tr><tr><td>P. vulgaris</td><td>0.1369 ± 0.0010</td><td>15.32 ± 0.18</td><td>19.93 ± 0.09</td><td>5.07 ± 0.04</td></tr><tr><td>P</td><td>&lt; 0.0001</td><td>0.0007</td><td>0.0013</td><td>&lt; 0.0001</td></tr></tbody></table>
information for forecasting and developing pest management strategies (Sedaratian *et al.* 2011; Segura-Martinez *et al.* 2023).

In summary, we show that *T. phaselus* had shorter developmental period, higher egg-to-adult survival and produced more daily and lifetime number of eggs on *G. max* than on *P. vulgaris*. Furthermore, females fed on *G. max* had significantly higher intrinsic rate of increase (*r*<sub>m</sub>) and net reproductive rate (*Ro*) and shorter generation time (*T*) and doubling time (*Dt*). These results suggest that *G. max* is more susceptible to *T. phaselus* attack. Considering that *G. max* and *P. vulgaris* are favorable host plants for *T. phaselus*, the increasing planting area of these beans particularly soybeans may facilitate the occurrence of *T. phaselus*.

## Acknowledgments

We thank Professor Tian-Ci Yi for identification of this spider mite to species. We also thank two anonymous reviewers and the editor, Dr. Dejan Marčić, for their constructive comments and suggestions, which have significantly improved the paper. The work reported here was supported by the Key Scientific Research Programs of Universities in Anhui Province (2022AH051052; 2024AH051125; 2024AH051080; 2024AH040170), and the Provincial Graduate Education Quality Engineering Project of Anhui Province (2023zybj021; 2024yjshhsfkc040).

## ORCID

Chen Chen https://orcid.org/0009-0002-0949-5874

Peng Zhou https://orcid.org/0009-0006-6347-4790

## References

Adango E., Onzo A., Hanna R., Atachi P., James B. 2006. Comparative demography of the spider mite, *Tetranychus ludeni*, on two host plants in West Africa. J. Insect Sci., 6: 1-9. https://doi.org/10.1673/031.006.4901

Alcázar-Valle M., Lugo-Cervantes E., Mojica L., Morales-Hernández N., Reyes-Ramírez H., Enríquez-Vara J.N., García-Morales S. 2020. Bioactive compounds, antioxidant activity, and antinutritional content of legumes: a comparison between four *Phaseolus* species. Molecules, 25: 3528. https://doi.org/10.3390/molecules25153528

Archontoulis S.V., Miguez F.E. 2015. Nonlinear regression models and applications in agricultural research. Agron. J., 107: 786-798. https://doi.org/10.2134/agronj2012.0506

Breukel L.M., Post A. 1959. The influence of the manurial treatment of orchards on the population density of *Metatetranychus ulmi* (Koch) (Acari, Tetranychidae). Entomol. Exp. Appl., 2: 38-47. https://doi.org/10.1111/j.1570-7458.1959.tb02095.x

Dahirel M., Masier S., Renault D., Bonte D. 2019. The distinct phenotypic signatures of dispersal and stress in an arthropod model: from physiology to life history. J. Exp. Biol., 222: jeb203596. https://doi.org/10.1242/jeb.203596

de Lima R.P., Bezerra M.M., de Moraes G.J., Furtado I. 2017. Life table of the red spider mite *Tetranychus bastosi* (Acari: Tetranychidae) on different host plants. Acarologia, 57: 601-605. https://doi.org/10.24349/acarologia/20174177

de Lima R.P., Guzzo É.C., Breda M.O. 2022. Biological performance of *Tetranychus bastosi* (Acari: Tetranychidae) on different hosts. Acarologia, 62: 1210-1216. https://doi.org/10.24349/k2gx-swz0

Ehara S. 1960. On some Japanese tetranychid mites of economic importance. Jpn. J. Appl. Entomol. Zool., 4: 234-241. https://doi.org/10.1303/jjaez.4.234

FAO. Food and Agriculture Organization of the United Nations: Crops and livestock products [Internet] [assessed on 04 March 2025]. Available from: https://www.fao.org/faostat/en/#data/QCL/visualize

Golizadeh A., Ghavidel S., Razmjou J., Fathi S.A.A., Hassanpour M. 2017. Comparative life table analysis of *Tetranychus urticae* Koch (Acari: Tetranychidae) on ten rose cultivars. Acarologia, 57: 607-616. https://doi.org/10.1111/j.1744-7917.2010.01379.x

Gotoh T., Moriya D., Nachman G. 2015. Development and reproduction of five *Tetranychus* species (Acari: Tetranychidae): Do they all have the potential to become major pests? Exp. Appl. Acarol., 66: 453-79. https://doi.org/10.1007/s10493-015-9919-y

Ito K., Fukuda T., Arakawa R. 2017. Host plant and field density of *Tetranychus phaselus* Ehara (Acari: Tetranychidae). J. Acarol. Soc. Jpn., 26: 13-24. https://doi.org/10.2300/acar26.13
Jervis M.A., Copland M.J.W. & Harvey J.A. 2005. The life-cycle. In: Jervis M.A. (Ed). Insects as natural enemies: a practical perspective. Dordrecht: Springer. p. 73-165. https://doi.org/10.1007/1-4020-2625-0_2

Jin P.-Y., Tian L., Chen L., Hong X.-Y. 2018. Spider mites of agricultural importance in China, with focus on species composition during the last decade (2008-2017). Syst. Appl. Acarol., 23: 2087-2098. https://doi.org/10.11158/saa.23.11.1

Kasap İ. 2004. Effect of apple cultivar and of temperature on the biology and life table parameters of the twospotted spider mite *Tetranychus urticae*. Phytoparasitica, 32: 73-82. https://doi.org/10.1007/BF02980863

Li G.-Y., Zhang Z.-Q. 2018. Does size matter? Fecundity and longevity of spider mites (Tetranychus urticae) in relation to mating and food availability. Syst. Appl. Acarol., 23: 1796-1808. https://doi.org/10.11158/saa.23.9.6

Li S., Chen J., Hao X., Ji X., Zhu Y., Chen X., Yao Y. 2024. A systematic review of black soybean (Glycine max (L.) Merr.): nutritional composition, bioactive compounds, health benefits, and processing to application. Food Front., 5: 1188-1211. https://doi.org/10.1002/ftf2.376

Migeon A., Dorkeld F. 2011. Spider Mites Web: a comprehensive database for the Tetranychidae. Trends in Acarology: 557-560. [Available from: https://www1.montpellier.inrae.fr/CBGP/spmweb/ Acessed 24 Sept 2024]. https://doi.org/10.1007/978-90-481-9837-5_96

NBSC. National Bureau of Statistics of China: National data [Internet] [assessed on 04 Mar 2025].
Available from: https://data.stats.gov.cn/easyquery.htm?cn=C01

Osman M.A., Al Dhafar Z.M., Alqahtani A.M. 2019. Biological responses of the two-spotted spider mite, *Tetranychus urticae* to different host plant. Arch. Phytopathol. Plant Prot., 52: 1229-1238.
https://doi.org/10.1080/03235408.2019.1703299

Puspitarini R.D., Fernando I., Rachmawati R., Hadi M.S., Rizali A. 2021. Host plant variability affects the development and reproduction of *Tetranychus urticae*. Int. J. Acarol., 47: 381-386.
https://doi.org/10.1080/01647954.2021.1915377

Ristyadi D., He X.Z., Wang Q. 2022. Predator- and killed prey-induced fears bear significant cost to an invasive spider mite: implications in pest management. Pest Manag. Sci., 78: 5456-5462.
https://doi.org/10.1002/ps.7168

Santamaría M.E., Auger P., Martínez M., Migeon A., Castanera P., Díaz I., Navajas M., Ortego F. 2018. Host plant use by two distinct lineages of the tomato red spider mite, *Tetranychus evansi*, differing in their distribution range. J. Pest Sci., 91: 169-179. https://doi.org/10.1007/s10340-017-0852-1

Sedaratian A., Fathipour Y., Moharramipour S. 2011. Comparative life table analysis of *Tetranychus urticae* (Acari: Tetranychidae) on 14 soybean genotypes. Insect Sci., 18: 541-553. https://doi.org/10.1111/j.1744-7917.2010.01379.x

Segura-Martínez M.T.d.J., Ordaz-Silva S., Hernández-Juárez A., Heinz-Castro R.T.Q., Mora-Ravelo S.G., Chacón-Hernández J.C. 2023. Life table parameters of *Tetranychus merganser* Boudreaux (Acari: Tetranychidae) on five host plants. Insects, 14: 473. https://doi.org/10.3390/insects14050473

van den Boom C.E.M., van Beek T., Dicke M. 2003. Differences among plant species in acceptance by the spider mite *Tetranychus urticae* Koch. J. Appl. Entomol., 127: 177-183. https://doi.org/10.1046/j.1439-0418.2003.00726.x

Weerawansha N., Wang Q., He X.Z. 2022. Comparing the effects of social environments and life history traits on sex allocation in a haplodiploid spider mite. Syst. Appl. Acarol., 27: 2123 - 2130.
https://doi.org/10.11158/saa.27.10.20

Weerawansha N., Wang Q., He X.Z. 2023. Reproductive plasticity in response to the changing cluster size during the breeding period: a case study in a spider mite. Exp. Appl. Acarol., 91: 237-250.
https://doi.org/10.1007/s10493-023-00834-y

Wermelinger B., Oertli J.J., Baumgärtner J. 1991. Environmental factors affecting the life-tables of *Tetranychus urticae* (Acari: Tetranychidae) III. Host-plant nutrition. Exp. Appl. Acarol., 12: 259-274.
https://doi.org/10.1007/BF01193472

Wu J., Wang L., Fu J., Chen J., Wei S., Zhang S., Zhang J., Tang Y., Chen M., Zhu J. et al. 2020. Resequencing of 683 common bean genotypes identifies yield component trait associations across a north-south cline. Nat. Genet., 52: 118-125. https://doi.org/10.1038/s41588-019-0546-0

Zhao D.-X., Chen D.-S., Ge C., Gotoh T., Hong X.-Y. 2013. Multiple infections with Cardinium and two strains of wolbachia in the spider mite *Tetranychus phaselus* Ehara: revealing new forces driving the spread of wolbachia. PLoS One, 8: e54964. https://doi.org/10.1371/journal.pone.0054964

Zhou P., He X.Z., Chen C., Wang Q. 2021. Resource relocations in relation to dispersal in *Tetranychus ludeni* Zacher. Syst. Appl. Acarol., 26: 2018-2026. https://doi.org/10.11158/saa.26.11.3

Zhou P., He X.Z., Wang Q. 2018. Sons from virgin mothers produce more daughters in a haplodiploid mite. Syst. Appl. Acarol., 23: 1869-1878. https://doi.org/10.11158/saa.23.9.13

Zhou P., He X.Z., Chen C., Wang Q. 2024. Age and density of mated females affect dispersal strategies in spider mite *Tetranychus ludeni* Zacher. Insects, 15, 387. https://doi.org/10.3390/insects15060387