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Full Research Article

Biology, Morphometric Characteristics and Life Table Parameters of the Pulse Beetle, Callosobruchus maculatus (Fabricius) on Green Gram under Laboratory Conditions

S
S. Karthik1
L
L. Ramazeame1,*
N
N. Murugan1
N
N. Vinothini2
R
R. Athira1
1Department of Entomology, SRM College of Agricultural Sciences, SRM Institute of Science and Technology, Chengalpattu-603 201, Tamil Nadu, India.
2Department of Seed Science and Technology, SRM College of Agricultural Sciences, SRM Institute of Science and Technology, Chengalpattu-603 201, Tamil Nadu, India.

ABSTRACT

Background: The pulse beetle Callosobruchus maculatus (Fabricius) is a major pest of stored green gram, leading to significant losses in seed quantity and quality. This study looked at the developmental biology, morphological characteristics, reproduction, sex ratio and population growth of C. maculatus reared on green gram in controlled lab conditions (28 ± 2°C and 65-75% relative humidity.

Methods: The study was carried out under controlled laboratory conditions using a standardized culture of C. maculatus maintained on green gram, where developmental stages, morphometric traits, reproductive parameters and life table data were recorded through systematic observations of individually reared cohorts.

Results: The eggs hatched within 5 to 7 days. The larval stage lasted for 15 to 22 days followed by the prepupal and pupal stages, which lasted 1 to 2 days and 5 to 8 days, respectively. Adult males lived between 8 and 13 days, while females lived for 10 to 15 days. Generally, females lived longer. The average fecundity was 76.8 eggs per female and the emerging population had a female-biased sex ratio of 1:1.3. Size analysis showed a steady increase in body size across successive instars and clear differences between male and female adults. Life-table analysis indicated that the highest mortality occurred during the transition from the egg stage to the early larval stage, accounting for about 19% mortality. Life expectancy declined progressively with age, decreasing from 23.30 days at egg hatch to 2.50 days by 36-40 days. Population growth parameters showed a net reproductive rate (R0) of 9.52, an intrinsic rate of increase (rm) of 0.0895 and a finite rate of increase (λ) of 1.09, suggesting a strong potential for population growth on this host. Overall, the findings indicate that green gram provides favourable conditions for the development and multiplication of C. maculatus, emphasizing the need for effective pest management strategies after harvest.

INTRODUCTION

Green gram [Vigna radiata (L.) Wilczek] is one of the pulse crops that is extensively cultivated in tropical and subtropical zones and is believed to be nutrient-rich, takes less time to grow and fits well in any agricultural conditions. In countries such as India, green gram contributes significantly to food and nutritional security and serves as an important source of plant proteins, vitamins and minerals (Ghosh et al., 2022; Sharma et al., 2017). However, bruchid infestation tends to damage the prolonged storage of green gram, which leads to substantial qualitative and quantitative losses (Swella and Mushobozy, 2009; Kazemi et al., 2009).
       
Among the various storage pests, the pulse beetle C. maculatus has been observed to be one of the most devastating insects of stored legumes (Malaikozhundan and Raj, 2012; Rahman et al., 2022). This pest is widely known as a major storage insect that causes huge economic losses in legumes due to its rapid multiplication and internal feeding behaviour (Kalpna et al., 2022; Hamzavi et al., 2022; Bidar et al., 2021). Adult females always deposit eggs on the seed surface and the young ones emerge, bore into the seed and develop within it. It reduces the weight of the seed, its germination potential and its nutritional and market quality due to such feeding behaviour (Sharma et al., 2017; Asra et al., 2025). Its reproduction rate is high, the pest has fast life cycle and its ability to adapt to other host legumes makes it tough to be managed under ordinary storage conditions (Rahman et al., 2022; Hamzavi et al., 2022; Kazemi et al., 2009; Bidar et al., 2021). This pest is widely distributed and exhibits high reproductive potential and rapid development, leading to severe losses in stored legumes (Garima et al., 2021; Hamzavi et al., 2022; Bidar et al., 2021).
       
Understanding the morphometry, biology and population dynamics of C. maculatus is critical for the development of effective management and storage strategies. The morphometric differentiation assists in the separation of the developmental stages and defining the sexual dimorphism, which is beneficial in laboratory research as well as in population studies (Singh and Swami, 2024; Jui et al., 2022; Premkumari et al., 2023). Some biological parameters, such as development period, fecundity and survival, also depend on the type of host and the environment (Rahman et al., 2022; Ahmady et al., 2016; Kafom et al., 2017; Salunkhe and Gaikwad, 2023). Moreover, in-depth information about the mortality, reproductive output, or possible population growth can be found through the life table analysis that has been successfully utilized and widely applied in studies involving stored-product insects (Chakraborty and Mondal, 2015; Mishra et al., 2023; Lekshmi et al., 2023).
       
Although several studies have examined the biology and morphometrics of C. maculatus on different legume hosts (Augustine and Balikai, 2019; Bidar et al., 2021; Singh and Swami, 2024; Salunkhe and Gaikwad, 2023), information on its development and population dynamics on green gram under controlled laboratory conditions remains limited (Thakur and Pathania, 2013; Singh et al., 2021).Therefore, the current study aimed to investigate the developmental biology, morphometric traits, fecundity and life-table parameters of C. maculatus on green gram under controlled environment.

MATERIALS AND METHODS

Insect culture and rearing
 
The experiment was conducted during 2025-2026 in the Entomology Laboratory at SRM College of Agricultural Sciences, under controlled conditions of 28±2oC and 65-75% RH (Hamzavi et al., 2022; Ahmady et al., 2016). The green gram seeds were cleaned manually and dried under sunlight for 6-8 hours and then sorted in airtight containers to maintain quality (Swella and Mushobozy, 2009; Kazemi et al., 2009).
       
A stock culture of C. maculatus was established using adults emerging from naturally infested seeds collected from local markets and maintained for three generations to obtain a uniform population (Malaikozhundan and Raj, 2012; Asra et al., 2025). For culture initiation,100 g of sun-dried seeds was placed in each container and 10 newly bred adults were introduced for oviposition. The fresh eggs (0-24 h old) were collected and the seeds with a single egg were taken to perforated vials to avoid the overlapping of larvae (Jui et al., 2022; Bhubaneshwari Devi and Victoria Devi, 2014).
       
The developmental stages were monitored daily. Size and morphological characters were used to identify each larval instar stage (Singh and Swami, 2024; Jui et al., 2022) and thoracic width, abdominal shape and pygidia were used to identify the sex of adults (Premkumari et al., 2023).

Morphometric observations
 
Morphometric measurements of immature stages and adults were taken using a Leica stereo microscope. Parameters recorded included body length, body breadth, head capsule dimensions, antennal length, elytral measurements, pygidium characteristics, pronotum measurements and total body length and width (Fig 1). Ten individuals from each developmental stage were measured and mean ± SD values were calculated following the documented procedures (Singh and Swami, 2024; Jui et al., 2022).

Fig 1: Major external morphological structures of Callosobruchus maculatus.


 
Biological parameters
 
Fecundity and reproductive observations were carried out by pairing newly emerged adults in small plastic containers containing 100 g of green gram seeds.Each plastic container was filled with 100 g of green gram seeds, into which 10 pairs of newly emerged C. maculatus adults were introduced to initiate oviposition. Data recorded included the pre-oviposition period, oviposition period, post-oviposition period, egg production per female, egg hatchability and adult longevity (Malaikozhundan and Raj, 2012; Asra et al., 2025). Survival and sex ratio were assessed using a cohort of 100 eggs and the number of individuals reaching each developmental stage was recorded (Kazemi et al., 2009; Rahman et al., 2022).
 
Life table
 
Age-specific and female-based life tables were prepared using a cohort of 100 eggs (Bidar et al., 2021; Chakraborty and Mondal, 2015; Lekshmi et al., 2023). Daily observations provided data on survivorship (lx), number dying (dx), mortality rate (qx) and age-specific fecundity (mx). Life expectancy (ex) and total remaining life-days (Tx) were calculated using standard demographic procedures (Lekshmi et al., 2023). Population growth parameters, including the net reproductive rate (R0), intrinsic rate of increase (rm), finite rate of increase (λ) and corrected generation time (T), were derived from cumulative lxmx and x·lxmx values (Bidar et al., 2021; Lekshmi et al., 2023).
 
Net reproductive rate (R0)
 
The net reproductive rate describes the total number of female offspring that a female contributes to the next generation over her lifetime.
R0= ∑lxmx
 
Mean generation time (Tc)
 
Mean generation time represents the average age at which females give birth to the next generation.

 
Intrinsic rate of increase (rm)
 
The intrinsic rate of increase is a measure of the population’s potential growth under ideal conditions.
 
∑e-rm.x.lxmx = 1

Corrected generation time (T)
 
Corrected generation time expresses the average time required for the population to grow by a factor of at the intrinsic rate of increase.

 
Finite rate of increase (λ)
 
The finite rate of increase shows the number of times the population multiplies per female per day.
 
λ = antilog (rm)
 
Life expectancy (ex)
 
Life expectancy at a given age represents the mean number of days an individual is expected to live beyond that age.

 
Age-class survivorship (Lx)
 
Survivorship within an age interval is expressed as Lx, which is the average number of individuals alive between ages x and x+1.

 
Mortality rate (qx)
 
Mortality rate indicates the proportion of individuals dying within a specific age interval.

 
Total remaining life-days (Tx)
 
Tx denotes the cumulative number of life-days lived by all individuals beyond age x. It is calculated as:

RESULTS AND DISCUSSION

Developmental biology
 
The biological characters of C. maculatus on green gram are presented in Table 1. Eggs hatched in 5.0-7.0 days (5.8 ±0.65 days) with a high hatching percentage of 82.0-94.0% (88.1±3.60%), indicating a favourable microenvironment for embryonic development. Similar observations on egg incubation period and hatchability have been reported in pulse beetles reared on legumes (Bhubaneshwari Devi and Victoria Devi, 2014; Thakur and Pathania, 2013; Augustine and Balikai, 2019; Rahman et al., 2022).

Table 1: Biology of pulse beetle, Callosobruchus maculatus on green gram under laboratory conditions.


       
The larval instars showed clear variability in developmental duration. The first instar lasted 5.0-7.0 days (6.1±0.65 days), followed by 3.0-4.0 days (3.6±0.48 days), 3.0-5.0 days (4.1±0.54 days) and 4.0-6.0 days (4.7±0.60 days) for the second, third and fourth instars, respectively, with the total larval period ranging from 15.0-22.0 days (18.4 ±1.32 days). The prepupal stage lasted 1.0-2.0 days (1.5± 0.35 days) and the pupal period 5.0-8.0 days (6.6±0.72 days) (Fig 2). Comparable developmental durations have been reported previously (Jui et al., 2022; Thakur and Pathania, 2013; Singh et al., 2021).

Fig 2: Developmental stages of Callosobruchus maculatus on green gram.


       
Adult lifespan differed by sex, with females showing greater longevity. Males lived 8.0-13.0 days (10.6±0.90 days) and females 10.0-15.0 days (12.2±1.12 days). The total life cycle lasted 22.0-34.0 days (28.1±1.95 days) in males and 25.0–38.0 days (31.2±2.18 days) in females. Similar findings on adult longevity and life cycle duration have been reported earlier (Malaikozhundan and Raj, 2012; Bidar et al., 2021; Asra et al., 2025).
 
Morphometric characteristics
 
Morphometric analysis of C. maculatus on green gram showed a gradual increase in body size from egg to adult (Table 2 and 3), confirming that morphometric characters can reliably identify developmental stages. Mean egg length ranged from 0.36-0.57 mm (0.46±0.05 mm) and breadth from 0.22-0.49 mm (0.35±0.07 mm), which agrees with earlier reports (Bhubaneshwari Devi and Victoria Devi, 2014; Hosamani et al., 2018).

Table 2: Morphometric characters of different immature stages and adults of pulse beetle, Callosobruchus maculatus.



Table 3: Comparative morphometric measurements of male and female pulse beetle, Callosobruchus maculatus.


       
Larval body length increased steadily from 0.68±0.04 mm in the first instar to 4.01±0.18 mm in the fourth instar, while breadth increased from 0.27±0.03 mm to 2.11±0.10 mm. Similar trends in larval growth have been documented previously (Singh and Swami, 2024; Mishra et al., 2023).
       
Adult morphometrics revealed clear sexual size dimorphism. Females were larger than males in most morphometric traits, whereas males possessed relatively longer antennae (Table 3). These observations are in close agreement with earlier findings on C. maculatus morphology (Premkumari et al., 2023; Kafom et al., 2017; Singh and Swami, 2024).
 
Fecundity, hatchability and sex ratio
 
Fecundity and oviposition behaviour are presented in Table 4. The pre-oviposition period ranged from 4.0-7.0 days (5.4± 1.05 days), oviposition period from 5.0-10.0 days (7.3±1.10 days) and post-oviposition period from 1.0-3.0 days (2.1± 0.45 days). Mean fecundity was 76.8±4.10 eggs per female and egg hatchability was 88.1±3.60%.

Table 4: Fecundity and oviposition behaviour of pulse beetle, Callosobruchus maculatus.


       
The high fecundity and hatchability observed in the present study indicate that green gram is a favourable host for the development and reproduction of C. maculatus. Similar observations have been reported previously (Malaikozhundan and Raj, 2012; Sharma et al., 2017; Rahman et al., 2022; Asra et al., 2025).
       
The sex ratio was female-biased (1:1.3, male:female), favouring faster population increase under storage conditions. Similar female-biased sex ratios have been observed in earlier studies (Kazemi et al., 2009; Bidar et al., 2021).
 
Life table and survivorship
 
Age-specific and female-based life tables are presented in Table 5 and 7. Survival declined from 100% at day 0 to 81% at 6-10 days and to 8% at 36-40 days, with the highest mortality occurring during the egg and early larval stages. Similar mortality patterns have been reported in pulse beetle populations maintained on legumes (Chakraborty and Mondal, 2015; Mishra et al., 2023; Lekshmi et al., 2023).

Table 5: Age-specific life table of Callosobruchus maculatus on green gram.


       
Life expectancy at hatching was 23.30 days and declined progressively to 2.50 days at 36-40 days (Table 5), which is consistent with previous reports on survivorship and life expectancy in stored-product bruchids (Mishra et al., 2023; Lekshmi et al., 2023).
 
Population growth parameters
 
Population growth parameters are presented in Table 6 and 7. Female survival (lx) remained constant at 0.74 during the immature period (0-22 days) and subsequently declined with increasing age, reaching 0.05 by day 34. Age-specific fecundity (mx) reached a maximum of 3.10 female offspring per female at 23 days of age and declined progressively thereafter. Similar reproductive trends have been reported in life-table studies of C. maculatus on different legume hosts (Bidar et al., 2021; Hamzavi et al., 2022).

Table 6: Life table (for female) and age specific fecundity for pulse beetle, Callosobruchus maculatus.



Table 7: Mean length of generation, innate capacity of increase in numbers and finite rate of increase in numbers of pulse beetle, Callosobruchus maculatus.


       
The net reproductive rate (R0) was 9.52 female offspring per female, the corrected generation time (T) was 25.17 days, the intrinsic rate of increase (rm) was 0.0895 females/day and the finite rate of increase (λ) was 1.094. These values indicate a strong capacity for population growth on green gram and are comparable to those reported previously for C. maculatus reared on suitable legume hosts (Bidar et al., 2021; Hamzavi et al., 2022; Lekshmi et al., 2023).           

CONCLUSION

The present research article has clearly demonstrated that green gram is so suited as a host to the proliferation and multiplication of the Callosobruchus maculatus. All the biological stages of egg to adult were effectively achieved, egg hatchability was high, the larvae and pupal stages were short and the sex ratio was female-biased, which was favourable to a high rate of population increase. The morphometric measurements provided a distinct, clear difference between the developmental stages and sexes, which confirmed the effectiveness of these traits in accurately determining the stage and in laboratory studies. The characteristics of the life table, including the high net reproductive rate (R0=9.52), intrinsic rate of increase (rm= 0.0895) and the finite rate of increase (λ=1.094), indicate the high reproduction capacity and a rapid multiplication of the beetle on green gram. These results indicate how stored green gram can be subject to severe bruchid infestation and the importance of effective storage pest management.

ACKNOWLEDGEMENT

The present study was supported by the Department of Entomology, SRM College of Agricultural Sciences, SRM Institute of Science and Technology, Kattankulathur, Tamil Nadu, India. The authors are thankful to the department for providing the necessary facilities to carry out this research work.
 
Disclaimers
 
The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.
 
Informed consent
 
All insect rearing and experimental procedures were conducted in accordance with standard laboratory protocols. The study involved only invertebrate insects and did not require approval from an animal ethics committee. No vertebrate animals were used in this study.

CONFLICT OF INTEREST

The authors declare that there are no conflicts of interest regarding the publication of this article. No funding or sponsorship influenced the design of the study, data collection, analysis, decision to publish, or preparation of the manuscript.

REFERENCES


  1. Ahmady, A., Rahmatzai, N., Zainullah, H., Mousa, M. and Zaitoun, A. (2016). Effect of temperature on the biology and morphometric measurement of cowpea beetle, Callosobruchus maculatus Fabr. (Coleoptera: Chrysomelidae) in cowpea seed. International Journal of Entomology Research. 1(7): 5-9.

  2. Asra, A., Syahrani, W., Maharani, B., Atsariyah, L., Joshi, R., Brugman, E., Fernando, I. and Mario, M. (2025). Biology of Callosobruchus  maculatus (Fabricius) (Coleoptera: Chrysomelidae) on two commercial beans in Makassar, Indonesia. Kufa Journal for Agricultural Sciences. 17: 155-164. https:// doi.org/10.36077/kjas/2025/v17i4.18666

  3. Augustine, N. and Balikai, R.A. (2019). Biology of pulse beetle, Callosobruchus chinensis (Linnaeus) on cowpea variety DC-15. Journal of Entomology and Zoology Studies. 7(1): 513-516.

  4. Bhubaneshwari Devi, M. and Victoria Devi, N. (2014). Biology and morphometric measurement of cowpea weevil, Callosobruchus maculatus Fabr. (Coleoptera: Chrysomelidae) in green gram. Journal of Entomology and Zoology Studies. 2: 74-77.

  5. Bidar, F., Razmjou, J., Golizadeh, A., Fathi, S.A.A., Ebadollahi, A. and Naseri, B. (2021). Effect of different legume seeds on life table parameters of cowpea weevil, Callosobruchus maculatus (F.) (Coleoptera: Chrysomelidae). Journal of Stored Products Research. 90: 101755. https://doi.org/ 10.1016/j.jspr.2020.101755

  6. Chakraborty, S. and Mondal, P. (2015). Age-specific and female fecundity life table of Callosobruchus chinensis on green gram. International Journal of Pure and Applied Bioscience. 3: 284-291.

  7. Garima, G., Khan, R. and Seal, D. (2021). Cowpea weevil, Callosobruchus maculatus (Insecta: Coleoptera: Bruchidae): EENY-769/IN1338. EDIS. 2021. https://doi.org/10.32473/ edis-in1338-2021.

  8. Ghosh, S., Roy, A. and Kundagrami, S. (2022). Screening of mungbean [Vigna radiata (L.) Wilczek] genotypes against bruchid (Callosobruchus maculatus) attack to reduce postharvest losses. Legume Research. 45(8): 1019-1027. doi: 10.18805/LR-4354.

  9. Hamzavi, F., Naseri, B., Hassanpour, M., Razmjou, J. and Golizadeh, A. (2022). Biology and life table parameters of Callosobruchus maculatus (F.) on Vigna unguiculata (L.) Walp. fertilized with some mineral- and bio-fertilizers. Journal of Stored Products Research. 97: 101978. https://doi.org/10.1016/ j.jspr.2022.101978.

  10. Hosamani, G.B., Jagginavar, S.B. and Karabhantanal, S.S. (2018). Biology of pulse beetle Callosobruchus chinensis on different pulses. Journal of Entomology and Zoology Studies. 6: 1898-1900.

  11. Jui, H., Swapon, M.A., Sultana, M., Rahman, M. and Rahman. (2022). Biology and morphometrics of pulse beetle reared on mungbean. Seed Technology. 4: 88-98.

  12. Kafom, A., Sarr, A., Dia, C.A.K.M., Ndiaye, M., Ngom, D. and Thiaw, C. (2017). Morphometric characterization of the Senegalese population of cowpea weevil: Callosobruchus maculatus (Fabricius, 1775). International Journal of Current Advanced Research. 5(8): 1734-1745.

  13. Kalpna, Hajam, Y. and Kumar, R. (2022). Management of stored grain pest with special reference to Callosobruchus maculatus, a major pest of cowpea: A review. Heliyon. 8: e08703. https://doi.org/10.1016/j.heliyon.2021.e08703.

  14. Kazemi, F., Talebi, A., Fathipour, Y. and Farahani, S. (2009). A comparative study on the effect of four leguminous species on biological and population growth parameters of Callosobruchus maculatus (F.) (Col.: Bruchidae). Advances in Environmental Biology. 3: 226-232.

  15. Lekshmi, J. K., Kennedy, J. S., Murugan, M. and Malarvizhi. (2023). A comparative analysis of age-stage two sex life tables in distinct forms of Callosobruchus maculatus (F.): Insights into population dynamics and reproductive strategies. Journal of Advanced Zoology. 44: 213-224. https:// doi.org/10.17762/jaz.v44i2.259.

  16. Malaikozhundan, B. and Raj, S. (2012). A study on the developmental biology of Callosobruchus maculatus (Fabricius) in different pulses. Legume Research. 35(2): 159-163.

  17. Mishra, O., Singh, K., Haldhar, S., Sinha, B., Singh, N., Chiranjiv, Roy, P. and Ibrar, S. (2023). Biology and morphometrics of pulse beetle, Callosobruchus chinensis L. on chickpea. International Journal of Economic Plants. 10: 295-301. https://doi.org/10.23910/2/2023.4937a

  18. Premkumari, S., Meena, R.S., Chauhan, R., Keval, R. and Srivastava, C.P. (2023). Biology and morphometric description of pulse beetle, Callosobruchus maculatus (Fabricius) on chickpea. Journal of Entomological Research. 47: 587- 591. https://doi.org/10.5958/0974-4576.2023.00108.1

  19. Rahman, M., Nesa, M., Sultana, S., Ara, N. and Laz, R. (2022). Studies on the life history traits of Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) reared in black gram (Vigna mungo L.). International Journal of Entomology Research. 7(4): 1-10.

  20. Salunkhe, D. and Gaikwad, S. (2023). Life stages of cowpea weevil Callosobruchus maculatus (F). Indian Journal of Entomology. 84(6): 1334-1337. https://doi.org/10.55446/IJE.2023.1099

  21. Sharma, R., Devi, R., Yadav, S. and Godara, P. (2017). Biology of pulse beetle, Callosobruchus maculatus (F.) and its response to botanicals in stored pigeonpea, Cajanus cajan (L.) grains. Legume Research. 41(6): 925-929. doi: 10.18805/LR-3807.

  22. Singh, G., Rana, B.S., Lal, J. and Sharma, K.C. (2021). Biology of pulse beetle [Callosobruchus chinensis (L.)] and its reaction against greengram [Vigna radiata (L.) R. Wilczek] varieties. Journal of Entomological Research. 45: 393- 398. https://doi.org/10.5958/0974-4576.2021.00062.1.

  23. Singh, N. and Swami, V. (2024). Morphological and morphometrical analysis of Callosobruchus maculatus (Coleoptera: Bruchidae) reared on green gram (Vigna radiata L.) and black gram (Vigna mungo L.). Journal of Entomology and Zoology Studies. 12: 122-127. https://doi.org/10.22271/j.ento. 2024.v12.i1b.9287.

  24. Swella, G. and Mushobozy, D. (2009). Comparative susceptibility of different legume seeds to infestation by cowpea bruchid Callosobruchus maculatus (F.) (Coleoptera: Chrysomelidae). Plant Protection Science. 45: 19-25. https://doi.org/ 10.17221/525-PPS.

  25. Thakur, A.K. and Pathania, M. (2013). Biology of pulse beetle (Callosobruchus chinensis) and its management through plant products on black gram (Vigna mungo). Science, Technology and Arts Research Journal. 2: 18. https:// doi.org/10.4314/star.v2i1.98838.
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    Full Research Article

    Biology, Morphometric Characteristics and Life Table Parameters of the Pulse Beetle, Callosobruchus maculatus (Fabricius) on Green Gram under Laboratory Conditions

    S
    S. Karthik1
    L
    L. Ramazeame1,*
    N
    N. Murugan1
    N
    N. Vinothini2
    R
    R. Athira1
    1Department of Entomology, SRM College of Agricultural Sciences, SRM Institute of Science and Technology, Chengalpattu-603 201, Tamil Nadu, India.
    2Department of Seed Science and Technology, SRM College of Agricultural Sciences, SRM Institute of Science and Technology, Chengalpattu-603 201, Tamil Nadu, India.

    ABSTRACT

    Background: The pulse beetle Callosobruchus maculatus (Fabricius) is a major pest of stored green gram, leading to significant losses in seed quantity and quality. This study looked at the developmental biology, morphological characteristics, reproduction, sex ratio and population growth of C. maculatus reared on green gram in controlled lab conditions (28 ± 2°C and 65-75% relative humidity.

    Methods: The study was carried out under controlled laboratory conditions using a standardized culture of C. maculatus maintained on green gram, where developmental stages, morphometric traits, reproductive parameters and life table data were recorded through systematic observations of individually reared cohorts.

    Results: The eggs hatched within 5 to 7 days. The larval stage lasted for 15 to 22 days followed by the prepupal and pupal stages, which lasted 1 to 2 days and 5 to 8 days, respectively. Adult males lived between 8 and 13 days, while females lived for 10 to 15 days. Generally, females lived longer. The average fecundity was 76.8 eggs per female and the emerging population had a female-biased sex ratio of 1:1.3. Size analysis showed a steady increase in body size across successive instars and clear differences between male and female adults. Life-table analysis indicated that the highest mortality occurred during the transition from the egg stage to the early larval stage, accounting for about 19% mortality. Life expectancy declined progressively with age, decreasing from 23.30 days at egg hatch to 2.50 days by 36-40 days. Population growth parameters showed a net reproductive rate (R0) of 9.52, an intrinsic rate of increase (rm) of 0.0895 and a finite rate of increase (λ) of 1.09, suggesting a strong potential for population growth on this host. Overall, the findings indicate that green gram provides favourable conditions for the development and multiplication of C. maculatus, emphasizing the need for effective pest management strategies after harvest.

    INTRODUCTION

    Green gram [Vigna radiata (L.) Wilczek] is one of the pulse crops that is extensively cultivated in tropical and subtropical zones and is believed to be nutrient-rich, takes less time to grow and fits well in any agricultural conditions. In countries such as India, green gram contributes significantly to food and nutritional security and serves as an important source of plant proteins, vitamins and minerals (Ghosh et al., 2022; Sharma et al., 2017). However, bruchid infestation tends to damage the prolonged storage of green gram, which leads to substantial qualitative and quantitative losses (Swella and Mushobozy, 2009; Kazemi et al., 2009).
           
    Among the various storage pests, the pulse beetle C. maculatus has been observed to be one of the most devastating insects of stored legumes (Malaikozhundan and Raj, 2012; Rahman et al., 2022). This pest is widely known as a major storage insect that causes huge economic losses in legumes due to its rapid multiplication and internal feeding behaviour (Kalpna et al., 2022; Hamzavi et al., 2022; Bidar et al., 2021). Adult females always deposit eggs on the seed surface and the young ones emerge, bore into the seed and develop within it. It reduces the weight of the seed, its germination potential and its nutritional and market quality due to such feeding behaviour (Sharma et al., 2017; Asra et al., 2025). Its reproduction rate is high, the pest has fast life cycle and its ability to adapt to other host legumes makes it tough to be managed under ordinary storage conditions (Rahman et al., 2022; Hamzavi et al., 2022; Kazemi et al., 2009; Bidar et al., 2021). This pest is widely distributed and exhibits high reproductive potential and rapid development, leading to severe losses in stored legumes (Garima et al., 2021; Hamzavi et al., 2022; Bidar et al., 2021).
           
    Understanding the morphometry, biology and population dynamics of C. maculatus is critical for the development of effective management and storage strategies. The morphometric differentiation assists in the separation of the developmental stages and defining the sexual dimorphism, which is beneficial in laboratory research as well as in population studies (Singh and Swami, 2024; Jui et al., 2022; Premkumari et al., 2023). Some biological parameters, such as development period, fecundity and survival, also depend on the type of host and the environment (Rahman et al., 2022; Ahmady et al., 2016; Kafom et al., 2017; Salunkhe and Gaikwad, 2023). Moreover, in-depth information about the mortality, reproductive output, or possible population growth can be found through the life table analysis that has been successfully utilized and widely applied in studies involving stored-product insects (Chakraborty and Mondal, 2015; Mishra et al., 2023; Lekshmi et al., 2023).
           
    Although several studies have examined the biology and morphometrics of C. maculatus on different legume hosts (Augustine and Balikai, 2019; Bidar et al., 2021; Singh and Swami, 2024; Salunkhe and Gaikwad, 2023), information on its development and population dynamics on green gram under controlled laboratory conditions remains limited (Thakur and Pathania, 2013; Singh et al., 2021).Therefore, the current study aimed to investigate the developmental biology, morphometric traits, fecundity and life-table parameters of C. maculatus on green gram under controlled environment.

    MATERIALS AND METHODS

    Insect culture and rearing
     
    The experiment was conducted during 2025-2026 in the Entomology Laboratory at SRM College of Agricultural Sciences, under controlled conditions of 28±2oC and 65-75% RH (Hamzavi et al., 2022; Ahmady et al., 2016). The green gram seeds were cleaned manually and dried under sunlight for 6-8 hours and then sorted in airtight containers to maintain quality (Swella and Mushobozy, 2009; Kazemi et al., 2009).
           
    A stock culture of C. maculatus was established using adults emerging from naturally infested seeds collected from local markets and maintained for three generations to obtain a uniform population (Malaikozhundan and Raj, 2012; Asra et al., 2025). For culture initiation,100 g of sun-dried seeds was placed in each container and 10 newly bred adults were introduced for oviposition. The fresh eggs (0-24 h old) were collected and the seeds with a single egg were taken to perforated vials to avoid the overlapping of larvae (Jui et al., 2022; Bhubaneshwari Devi and Victoria Devi, 2014).
           
    The developmental stages were monitored daily. Size and morphological characters were used to identify each larval instar stage (Singh and Swami, 2024; Jui et al., 2022) and thoracic width, abdominal shape and pygidia were used to identify the sex of adults (Premkumari et al., 2023).

    Morphometric observations
     
    Morphometric measurements of immature stages and adults were taken using a Leica stereo microscope. Parameters recorded included body length, body breadth, head capsule dimensions, antennal length, elytral measurements, pygidium characteristics, pronotum measurements and total body length and width (Fig 1). Ten individuals from each developmental stage were measured and mean ± SD values were calculated following the documented procedures (Singh and Swami, 2024; Jui et al., 2022).

    Fig 1: Major external morphological structures of Callosobruchus maculatus.


     
    Biological parameters
     
    Fecundity and reproductive observations were carried out by pairing newly emerged adults in small plastic containers containing 100 g of green gram seeds.Each plastic container was filled with 100 g of green gram seeds, into which 10 pairs of newly emerged C. maculatus adults were introduced to initiate oviposition. Data recorded included the pre-oviposition period, oviposition period, post-oviposition period, egg production per female, egg hatchability and adult longevity (Malaikozhundan and Raj, 2012; Asra et al., 2025). Survival and sex ratio were assessed using a cohort of 100 eggs and the number of individuals reaching each developmental stage was recorded (Kazemi et al., 2009; Rahman et al., 2022).
     
    Life table
     
    Age-specific and female-based life tables were prepared using a cohort of 100 eggs (Bidar et al., 2021; Chakraborty and Mondal, 2015; Lekshmi et al., 2023). Daily observations provided data on survivorship (lx), number dying (dx), mortality rate (qx) and age-specific fecundity (mx). Life expectancy (ex) and total remaining life-days (Tx) were calculated using standard demographic procedures (Lekshmi et al., 2023). Population growth parameters, including the net reproductive rate (R0), intrinsic rate of increase (rm), finite rate of increase (λ) and corrected generation time (T), were derived from cumulative lxmx and x·lxmx values (Bidar et al., 2021; Lekshmi et al., 2023).
     
    Net reproductive rate (R0)
     
    The net reproductive rate describes the total number of female offspring that a female contributes to the next generation over her lifetime.
    R0= ∑lxmx
     
    Mean generation time (Tc)
     
    Mean generation time represents the average age at which females give birth to the next generation.

     
    Intrinsic rate of increase (rm)
     
    The intrinsic rate of increase is a measure of the population’s potential growth under ideal conditions.
     
    ∑e-rm.x.lxmx = 1

    Corrected generation time (T)
     
    Corrected generation time expresses the average time required for the population to grow by a factor of at the intrinsic rate of increase.

     
    Finite rate of increase (λ)
     
    The finite rate of increase shows the number of times the population multiplies per female per day.
     
    λ = antilog (rm)
     
    Life expectancy (ex)
     
    Life expectancy at a given age represents the mean number of days an individual is expected to live beyond that age.

     
    Age-class survivorship (Lx)
     
    Survivorship within an age interval is expressed as Lx, which is the average number of individuals alive between ages x and x+1.

     
    Mortality rate (qx)
     
    Mortality rate indicates the proportion of individuals dying within a specific age interval.

     
    Total remaining life-days (Tx)
     
    Tx denotes the cumulative number of life-days lived by all individuals beyond age x. It is calculated as:

    RESULTS AND DISCUSSION

    Developmental biology
     
    The biological characters of C. maculatus on green gram are presented in Table 1. Eggs hatched in 5.0-7.0 days (5.8 ±0.65 days) with a high hatching percentage of 82.0-94.0% (88.1±3.60%), indicating a favourable microenvironment for embryonic development. Similar observations on egg incubation period and hatchability have been reported in pulse beetles reared on legumes (Bhubaneshwari Devi and Victoria Devi, 2014; Thakur and Pathania, 2013; Augustine and Balikai, 2019; Rahman et al., 2022).

    Table 1: Biology of pulse beetle, Callosobruchus maculatus on green gram under laboratory conditions.


           
    The larval instars showed clear variability in developmental duration. The first instar lasted 5.0-7.0 days (6.1±0.65 days), followed by 3.0-4.0 days (3.6±0.48 days), 3.0-5.0 days (4.1±0.54 days) and 4.0-6.0 days (4.7±0.60 days) for the second, third and fourth instars, respectively, with the total larval period ranging from 15.0-22.0 days (18.4 ±1.32 days). The prepupal stage lasted 1.0-2.0 days (1.5± 0.35 days) and the pupal period 5.0-8.0 days (6.6±0.72 days) (Fig 2). Comparable developmental durations have been reported previously (Jui et al., 2022; Thakur and Pathania, 2013; Singh et al., 2021).

    Fig 2: Developmental stages of Callosobruchus maculatus on green gram.


           
    Adult lifespan differed by sex, with females showing greater longevity. Males lived 8.0-13.0 days (10.6±0.90 days) and females 10.0-15.0 days (12.2±1.12 days). The total life cycle lasted 22.0-34.0 days (28.1±1.95 days) in males and 25.0–38.0 days (31.2±2.18 days) in females. Similar findings on adult longevity and life cycle duration have been reported earlier (Malaikozhundan and Raj, 2012; Bidar et al., 2021; Asra et al., 2025).
     
    Morphometric characteristics
     
    Morphometric analysis of C. maculatus on green gram showed a gradual increase in body size from egg to adult (Table 2 and 3), confirming that morphometric characters can reliably identify developmental stages. Mean egg length ranged from 0.36-0.57 mm (0.46±0.05 mm) and breadth from 0.22-0.49 mm (0.35±0.07 mm), which agrees with earlier reports (Bhubaneshwari Devi and Victoria Devi, 2014; Hosamani et al., 2018).

    Table 2: Morphometric characters of different immature stages and adults of pulse beetle, Callosobruchus maculatus.



    Table 3: Comparative morphometric measurements of male and female pulse beetle, Callosobruchus maculatus.


           
    Larval body length increased steadily from 0.68±0.04 mm in the first instar to 4.01±0.18 mm in the fourth instar, while breadth increased from 0.27±0.03 mm to 2.11±0.10 mm. Similar trends in larval growth have been documented previously (Singh and Swami, 2024; Mishra et al., 2023).
           
    Adult morphometrics revealed clear sexual size dimorphism. Females were larger than males in most morphometric traits, whereas males possessed relatively longer antennae (Table 3). These observations are in close agreement with earlier findings on C. maculatus morphology (Premkumari et al., 2023; Kafom et al., 2017; Singh and Swami, 2024).
     
    Fecundity, hatchability and sex ratio
     
    Fecundity and oviposition behaviour are presented in Table 4. The pre-oviposition period ranged from 4.0-7.0 days (5.4± 1.05 days), oviposition period from 5.0-10.0 days (7.3±1.10 days) and post-oviposition period from 1.0-3.0 days (2.1± 0.45 days). Mean fecundity was 76.8±4.10 eggs per female and egg hatchability was 88.1±3.60%.

    Table 4: Fecundity and oviposition behaviour of pulse beetle, Callosobruchus maculatus.


           
    The high fecundity and hatchability observed in the present study indicate that green gram is a favourable host for the development and reproduction of C. maculatus. Similar observations have been reported previously (Malaikozhundan and Raj, 2012; Sharma et al., 2017; Rahman et al., 2022; Asra et al., 2025).
           
    The sex ratio was female-biased (1:1.3, male:female), favouring faster population increase under storage conditions. Similar female-biased sex ratios have been observed in earlier studies (Kazemi et al., 2009; Bidar et al., 2021).
     
    Life table and survivorship
     
    Age-specific and female-based life tables are presented in Table 5 and 7. Survival declined from 100% at day 0 to 81% at 6-10 days and to 8% at 36-40 days, with the highest mortality occurring during the egg and early larval stages. Similar mortality patterns have been reported in pulse beetle populations maintained on legumes (Chakraborty and Mondal, 2015; Mishra et al., 2023; Lekshmi et al., 2023).

    Table 5: Age-specific life table of Callosobruchus maculatus on green gram.


           
    Life expectancy at hatching was 23.30 days and declined progressively to 2.50 days at 36-40 days (Table 5), which is consistent with previous reports on survivorship and life expectancy in stored-product bruchids (Mishra et al., 2023; Lekshmi et al., 2023).
     
    Population growth parameters
     
    Population growth parameters are presented in Table 6 and 7. Female survival (lx) remained constant at 0.74 during the immature period (0-22 days) and subsequently declined with increasing age, reaching 0.05 by day 34. Age-specific fecundity (mx) reached a maximum of 3.10 female offspring per female at 23 days of age and declined progressively thereafter. Similar reproductive trends have been reported in life-table studies of C. maculatus on different legume hosts (Bidar et al., 2021; Hamzavi et al., 2022).

    Table 6: Life table (for female) and age specific fecundity for pulse beetle, Callosobruchus maculatus.



    Table 7: Mean length of generation, innate capacity of increase in numbers and finite rate of increase in numbers of pulse beetle, Callosobruchus maculatus.


           
    The net reproductive rate (R0) was 9.52 female offspring per female, the corrected generation time (T) was 25.17 days, the intrinsic rate of increase (rm) was 0.0895 females/day and the finite rate of increase (λ) was 1.094. These values indicate a strong capacity for population growth on green gram and are comparable to those reported previously for C. maculatus reared on suitable legume hosts (Bidar et al., 2021; Hamzavi et al., 2022; Lekshmi et al., 2023).           

    CONCLUSION

    The present research article has clearly demonstrated that green gram is so suited as a host to the proliferation and multiplication of the Callosobruchus maculatus. All the biological stages of egg to adult were effectively achieved, egg hatchability was high, the larvae and pupal stages were short and the sex ratio was female-biased, which was favourable to a high rate of population increase. The morphometric measurements provided a distinct, clear difference between the developmental stages and sexes, which confirmed the effectiveness of these traits in accurately determining the stage and in laboratory studies. The characteristics of the life table, including the high net reproductive rate (R0=9.52), intrinsic rate of increase (rm= 0.0895) and the finite rate of increase (λ=1.094), indicate the high reproduction capacity and a rapid multiplication of the beetle on green gram. These results indicate how stored green gram can be subject to severe bruchid infestation and the importance of effective storage pest management.

    ACKNOWLEDGEMENT

    The present study was supported by the Department of Entomology, SRM College of Agricultural Sciences, SRM Institute of Science and Technology, Kattankulathur, Tamil Nadu, India. The authors are thankful to the department for providing the necessary facilities to carry out this research work.
     
    Disclaimers
     
    The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.
     
    Informed consent
     
    All insect rearing and experimental procedures were conducted in accordance with standard laboratory protocols. The study involved only invertebrate insects and did not require approval from an animal ethics committee. No vertebrate animals were used in this study.

    CONFLICT OF INTEREST

    The authors declare that there are no conflicts of interest regarding the publication of this article. No funding or sponsorship influenced the design of the study, data collection, analysis, decision to publish, or preparation of the manuscript.

    REFERENCES


    1. Ahmady, A., Rahmatzai, N., Zainullah, H., Mousa, M. and Zaitoun, A. (2016). Effect of temperature on the biology and morphometric measurement of cowpea beetle, Callosobruchus maculatus Fabr. (Coleoptera: Chrysomelidae) in cowpea seed. International Journal of Entomology Research. 1(7): 5-9.

    2. Asra, A., Syahrani, W., Maharani, B., Atsariyah, L., Joshi, R., Brugman, E., Fernando, I. and Mario, M. (2025). Biology of Callosobruchus  maculatus (Fabricius) (Coleoptera: Chrysomelidae) on two commercial beans in Makassar, Indonesia. Kufa Journal for Agricultural Sciences. 17: 155-164. https:// doi.org/10.36077/kjas/2025/v17i4.18666

    3. Augustine, N. and Balikai, R.A. (2019). Biology of pulse beetle, Callosobruchus chinensis (Linnaeus) on cowpea variety DC-15. Journal of Entomology and Zoology Studies. 7(1): 513-516.

    4. Bhubaneshwari Devi, M. and Victoria Devi, N. (2014). Biology and morphometric measurement of cowpea weevil, Callosobruchus maculatus Fabr. (Coleoptera: Chrysomelidae) in green gram. Journal of Entomology and Zoology Studies. 2: 74-77.

    5. Bidar, F., Razmjou, J., Golizadeh, A., Fathi, S.A.A., Ebadollahi, A. and Naseri, B. (2021). Effect of different legume seeds on life table parameters of cowpea weevil, Callosobruchus maculatus (F.) (Coleoptera: Chrysomelidae). Journal of Stored Products Research. 90: 101755. https://doi.org/ 10.1016/j.jspr.2020.101755

    6. Chakraborty, S. and Mondal, P. (2015). Age-specific and female fecundity life table of Callosobruchus chinensis on green gram. International Journal of Pure and Applied Bioscience. 3: 284-291.

    7. Garima, G., Khan, R. and Seal, D. (2021). Cowpea weevil, Callosobruchus maculatus (Insecta: Coleoptera: Bruchidae): EENY-769/IN1338. EDIS. 2021. https://doi.org/10.32473/ edis-in1338-2021.

    8. Ghosh, S., Roy, A. and Kundagrami, S. (2022). Screening of mungbean [Vigna radiata (L.) Wilczek] genotypes against bruchid (Callosobruchus maculatus) attack to reduce postharvest losses. Legume Research. 45(8): 1019-1027. doi: 10.18805/LR-4354.

    9. Hamzavi, F., Naseri, B., Hassanpour, M., Razmjou, J. and Golizadeh, A. (2022). Biology and life table parameters of Callosobruchus maculatus (F.) on Vigna unguiculata (L.) Walp. fertilized with some mineral- and bio-fertilizers. Journal of Stored Products Research. 97: 101978. https://doi.org/10.1016/ j.jspr.2022.101978.

    10. Hosamani, G.B., Jagginavar, S.B. and Karabhantanal, S.S. (2018). Biology of pulse beetle Callosobruchus chinensis on different pulses. Journal of Entomology and Zoology Studies. 6: 1898-1900.

    11. Jui, H., Swapon, M.A., Sultana, M., Rahman, M. and Rahman. (2022). Biology and morphometrics of pulse beetle reared on mungbean. Seed Technology. 4: 88-98.

    12. Kafom, A., Sarr, A., Dia, C.A.K.M., Ndiaye, M., Ngom, D. and Thiaw, C. (2017). Morphometric characterization of the Senegalese population of cowpea weevil: Callosobruchus maculatus (Fabricius, 1775). International Journal of Current Advanced Research. 5(8): 1734-1745.

    13. Kalpna, Hajam, Y. and Kumar, R. (2022). Management of stored grain pest with special reference to Callosobruchus maculatus, a major pest of cowpea: A review. Heliyon. 8: e08703. https://doi.org/10.1016/j.heliyon.2021.e08703.

    14. Kazemi, F., Talebi, A., Fathipour, Y. and Farahani, S. (2009). A comparative study on the effect of four leguminous species on biological and population growth parameters of Callosobruchus maculatus (F.) (Col.: Bruchidae). Advances in Environmental Biology. 3: 226-232.

    15. Lekshmi, J. K., Kennedy, J. S., Murugan, M. and Malarvizhi. (2023). A comparative analysis of age-stage two sex life tables in distinct forms of Callosobruchus maculatus (F.): Insights into population dynamics and reproductive strategies. Journal of Advanced Zoology. 44: 213-224. https:// doi.org/10.17762/jaz.v44i2.259.

    16. Malaikozhundan, B. and Raj, S. (2012). A study on the developmental biology of Callosobruchus maculatus (Fabricius) in different pulses. Legume Research. 35(2): 159-163.

    17. Mishra, O., Singh, K., Haldhar, S., Sinha, B., Singh, N., Chiranjiv, Roy, P. and Ibrar, S. (2023). Biology and morphometrics of pulse beetle, Callosobruchus chinensis L. on chickpea. International Journal of Economic Plants. 10: 295-301. https://doi.org/10.23910/2/2023.4937a

    18. Premkumari, S., Meena, R.S., Chauhan, R., Keval, R. and Srivastava, C.P. (2023). Biology and morphometric description of pulse beetle, Callosobruchus maculatus (Fabricius) on chickpea. Journal of Entomological Research. 47: 587- 591. https://doi.org/10.5958/0974-4576.2023.00108.1

    19. Rahman, M., Nesa, M., Sultana, S., Ara, N. and Laz, R. (2022). Studies on the life history traits of Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) reared in black gram (Vigna mungo L.). International Journal of Entomology Research. 7(4): 1-10.

    20. Salunkhe, D. and Gaikwad, S. (2023). Life stages of cowpea weevil Callosobruchus maculatus (F). Indian Journal of Entomology. 84(6): 1334-1337. https://doi.org/10.55446/IJE.2023.1099

    21. Sharma, R., Devi, R., Yadav, S. and Godara, P. (2017). Biology of pulse beetle, Callosobruchus maculatus (F.) and its response to botanicals in stored pigeonpea, Cajanus cajan (L.) grains. Legume Research. 41(6): 925-929. doi: 10.18805/LR-3807.

    22. Singh, G., Rana, B.S., Lal, J. and Sharma, K.C. (2021). Biology of pulse beetle [Callosobruchus chinensis (L.)] and its reaction against greengram [Vigna radiata (L.) R. Wilczek] varieties. Journal of Entomological Research. 45: 393- 398. https://doi.org/10.5958/0974-4576.2021.00062.1.

    23. Singh, N. and Swami, V. (2024). Morphological and morphometrical analysis of Callosobruchus maculatus (Coleoptera: Bruchidae) reared on green gram (Vigna radiata L.) and black gram (Vigna mungo L.). Journal of Entomology and Zoology Studies. 12: 122-127. https://doi.org/10.22271/j.ento. 2024.v12.i1b.9287.

    24. Swella, G. and Mushobozy, D. (2009). Comparative susceptibility of different legume seeds to infestation by cowpea bruchid Callosobruchus maculatus (F.) (Coleoptera: Chrysomelidae). Plant Protection Science. 45: 19-25. https://doi.org/ 10.17221/525-PPS.

    25. Thakur, A.K. and Pathania, M. (2013). Biology of pulse beetle (Callosobruchus chinensis) and its management through plant products on black gram (Vigna mungo). Science, Technology and Arts Research Journal. 2: 18. https:// doi.org/10.4314/star.v2i1.98838.
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