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Abstract
Wallago attu (Bloch & Schneider, 1801), an economically valuable silurid catfish species, is extensively harvested from South Asian river systems. However, information on stock levels in the Indus River population in Pakistan remains limited. This study examined morphological differences among three geographically distinct populations of W. attu from the Chashma, Sukkur, and Taunsa Barrages, located at different elevations in the Indus River system. Thirty-four morphometric characters were recorded from 75 individuals (25 per location) using a landmark method, and multivariate, univariate, and allometric analyses were conducted. Multivariate analysis of variance showed significant differentiation in body shape among the three W. attu populations (Pillai’s trace= 1.49, F₍₂₆, ₁₂₂₎= 13.80, P<0.001). Thirteen of the 34 characters showed significant differences after False Discovery Rate correction, while total, fork, and standard lengths showed no significant differences among the three populations, meaning there were no significant differences in size among the three populations. However, they showed significant differences in body proportions, specifically head region and fin traits, such as post-orbital length (η²= 0.31) and head depth (η²= 0.25). Principal component analysis showed size-based overlap along the first axis (53.2%) but partial segregation along shape axes, with the Chashma group as the most distinct cluster. Cross-validated discriminant analysis classified 84% of individuals to their sites of origin. Allometric growth was isometric in the Sukkur and Taunsa Barrage samples. In contrast, non-isometry was common among the Chashma fish, and the length-weight relation showed negative allometry in the Chashma and Sukkur samples and isometry at the Taunsa Barrage. This study shows that the three barrage samples represent morphologically distinct phenotypic stocks, most likely influenced by differences between lentic and lotic habitats in the Indus River.
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The authors transfer the copyrights of their papers to the Iranian Society of Ichthyology. However, the information could be used in accordance with the Creative Commons licence (Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
References
- Ali, A., Shah, S.; Ali, A. & Haris, M. 2024. Fish diversity in different streams of Lower Swat, Pakistan. Journal of Wildlife and Ecology 8: 144-153.
- Altaf, M. 2026. Freshwater Fishes of Pakistan: Ecology, Distribution and Threats. Journal of Wildlife and Ecology. 10(SI-1).
- Bagenal, T.B. & Tesch, F.W. 1978. Age and growth. In T. B. Bagenal (Ed.), Methods for assessment of fish production in fresh waters (3rd ed., pp. 101-136). Blackwell Scientific Publications.
- Basharat, H.; Ghafoor, A.; Chavhan, A.; Zafar, M.; Abbas, K. & Parveen, J. 2016. Microsatellite markers revealed poor genetic structure of Wallago attu in Punjab, Pakistan. International Journal of Life Sciences 4(3): 385-393.
- Begg, G.A. & Waldman, J.R. 1999. A holistic approach to fish stock identification. Fisheries Research 43(1-3): 35-44.
- Benjamini, Y. & Hochberg, Y. 1995. Controlling the false discovery rate: A practical and powerful approach to multiple testing. Journal of the Royal Statistical Society: Series B (Methodological) 57(1): 289-300.
- Cadrin, S.X. 2000. Advances in morphometric identification of fishery stocks. Reviews in Fish Biology and Fisheries 10(1): 91-112.
- Franssen, N.R. 2011. Anthropogenic habitat alteration induces rapid morphological divergence in a native stream fish. Evolutionary Applications 4(6): 791-804.
- Grill, G.; Lehner, B.; Thieme, M.; Geenen, B.; Tickner, D.; Antonelli, F.; Babu, S.; Borrelli, P.; Cheng, L.; Crochetiere, H. & Ehalt Macedo, H. 2019. Mapping the world's free-flowing rivers. Nature 569(7755): 215-221.
- Haas, T.C.; Blum, M.J. & Heins, D.C. 2010. Morphological responses of a stream fish to water impoundment. Biology Letters 6(6): 803-806.
- Habib, S.S.; Naz, S. & Mehmood, R. 2020. Morphometric relationships of some specified species of the family Cyprinidae in Jinnah Barrage, Punjab, Pakistan. Journal of Bioresource Management 7(2).
- Hetzel, C. & Forsythe, P. 2023. Phenotypic plasticity of a generalist fish species resident to lotic environments: Insights from the Great Lakes region. Ecology and Evolution 13(11): e10715.
- Imran, H., Zohra, K. & Osmany, H. 2025. Stomach analysis of the Milk shark, Rhizoprionodon acutus (Ruppell, 1837) in the Coastal Waters of the Pakistan, Northern Arabian Sea. Journal of Wildlife and Ecology 9: 268-281.
- Kalsoom, B.; Mishaal, B.; Sardar, T.; Nasreen, Z.; Kalsoom, S. & Sharif, A. 2021. Water quality parameters and fish diversity at Chashma Barrage, Mianwali, Punjab, Pakistan. RADS Journal of Biological Research & Applied Sciences 12(2): 135-145.
- Kelley, J.L.; Davies, P.M.; Collin, S.P. & Grierson, P.F. 2017. Morphological plasticity in a native freshwater fish from semiarid Australia in response to variable water flows. Ecology and Evolution 7(16): 6595-6605.
- Klingenberg, C.P. 2022. Methods for studying allometry in geometric morphometrics: A comparison of performance. Evolutionary Ecology 36(4): 439-470.
- Kumar, A.; Negi, N.; Yadav, N.; Badola, R.; Hussain, S.A. & Gupta, S.K. 2024. Genetic diversity, spatial connectivity, and population structure of Asian silurid catfish Wallago attu (Bloch and Schneider, 1801) in the Ganga River System: Insights from mitochondrial DNA analysis. Molecular Biology Reports 51(1): Article 378.
- Kumar, A.; Negi, N.; Yadav, N.; Badola, R.; Hussain, S.A. & Gupta, S.K. 2024. Genetic diversity, spatial connectivity, and population structure of Asian silurid catfish Wallago attu (Bloch and Schneider, 1801) in the Ganga River System: Insights from mitochondrial DNA analysis. Molecular Biology Reports 51, Article 378.
- Kumar, G.; Kashyap, A. & Serajuddin, M. 2023. Assessment of morphometric variations among the populations of Asian sheat catfish Wallago attu (Siluridae) from five Indian rivers. Inland Water Biology 16(6): 1026-1037.
- Kwikiriza, G.; Yegon, M.J.; Byamugisha, N.; Beingana, A.; Atukwatse, F.; Barekye, A. & Meimberg, H. 2023. Morphometric variations of Nile tilapia (Oreochromis niloticus) (Linnaeus, 1758) local strains collected from different fish farms in South Western Highland Agro-Ecological Zone, Uganda: Screening strains for aquaculture. Fishes 8(4): 217.
- Langerhans, R.B. 2008. Predictability of phenotypic differentiation across flow regimes in fishes. Integrative and Comparative Biology 48(6): 750-768.
- Langerhans, R.B. & Reznick, D.N. 2010. Ecology and evolution of swimming performance in fishes: Predicting evolution with biomechanics. In P. Domenici & B. G. Kapoor (Eds.), Fish locomotion: An eco-ethological perspective (pp. 200-248). Science Publishers.
- Le Cren, E.D. 1951. The length–weight relationship and seasonal cycle in gonad weight and condition in the perch (Perca fluviatilis). Journal of Animal Ecology 20(2): 201-219.
- Malik, D.S.; Sharma, A.K.; Sharma, A.K.; Thakur, R. & Sharma, M. 2020. A review on impact of water pollution on freshwater fish species and their aquatic environment. In Advances in Environmental Pollution Management: Wastewater Impacts and Treatment Technologies 1: 10-28.
- Mouludi-Saleh, A.; Keivany, Y. & Jalali, S.A.H. 2018. Comparison of meristic traits in Transcaucasian chub (Squalius turcicus De Filippi, 1865) from Caspian Sea basin. International Journal of Aquatic Biology 6(1): 8-14.
- Mirza, M. & Sandhu, A. 2007. Fishes of the Punjab, Pakistan. Lahore, Pakistan: Polymer Publications.
- Muhammad, N. & Zain-ul-Abideen, S.M. 2022. Diversity of fishes of head Marala, Punjab, Pakistan. Journal of Wildlife and Ecology 6(2): 58-63.
- Muhammad, N.; Rasool, F. & Imran, M. 2026. Mitochondrial ATPase 6/8-Based Assessment of Genetic Diversity and Population Structure in Cirrhinus mrigala Across the Indus River System, Pakistan. Journal of Wildlife and Ecology 10(SI-1).
- Muhammad, N.; Altaf, M.A.; Abbasi, M.; Khan, A.M. & Iqbal, K.J. 2023. From water to Remedy: Fishes as Ethnopharmacological Resources along the River Ravi, Punjab, Pakistan. Journal of Wildlife and Ecology 7(3): 93-108.
- Muhammad, N.; Imran, M. & Abideen, S. 2024. Exploring the fish diversity along the River Ravi in Punjab, Pakistan. Journal of Wildlife and Ecology 8: 74-81.
- Naseer, A.; Imran, M.; Khan, A.M. & Ammar, M. 2023. A comparative morphologic and craniometric analysis of grass carp (Ctenopharyngodon idella) population from Punjab, Pakistan. Journal of Xi’an Shiyou University 19(4): 48-62.
- Ng, H.H.; de Alwis Goonatilake, S.; Fernado, M. & Kotagama, O. 2020. Wallago attu. The IUCN Red List of Threatened Species. International Union for Conservation of Nature.
- Osmany, H.B.; Zohra, K. & Imran, H. 2026. Seasonal Variation in Feeding Habits of the Hardnose Shark (Carcharhinus macloti) and Comparison of Diet with Other Demersal Sharks in the Northern Arabian Sea of Pakistan. Journal of Wildlife and Ecology 10(SI-1).
- R Core Team. 2023. R: A language and environment for statistical computing. R Foundation for Statistical Computing.
- Rawat, S.; Benakappa, S.; Kumar, J.; Naik, A.S.K.; Pandey, G. & Pema, C.W. 2017. Identification of fish stocks based on truss morphometric: A review. Journal of Fisheries and Life Sciences 2(1): 9-14.
- Reist, J. D. 1985. An empirical evaluation of several univariate methods that adjust for size variation in morphometric data. Canadian Journal of Zoology 63(6): 1429-1439.
- Rohlf, F.J. 2015. The tps series of software. Hystrix, the Italian Journal of Mammalogy 26(1): 9-12.
- Secer, B.; Mouludi-Saleh, A.; Eagderi, S.; Poorbagher, H.; Cicek, E. & Sungur, S. 2022. Phenotypic plasticity of angora loach, Oxynoemacheilus angorae (steindachner, 1897) in inland waters of turkey. Iranian Journal of Science and Technology, Transactions A: Science 46(5): 1317-1326.
- Sajjad, A.; Jabeen, F.; Ali, M. & Zafar, S. 2023. DNA barcoding and phylogenetics of Wallago attu using mitochondrial COI gene from the River Indus. Journal of King Saud University–Science 35(6): 102725.
- Shabir, M., Bano, N. & Majeed, U. 2025. Effect of Fermented Moringa (Moringa oleifera) Leaves as a Feed Ingredient in the Diet of Rohu (Labeo rohita) Fish. Journal of Wildlife and Ecology 9(1): 31-42.
- Sherzada, S.; Hussain, N.; Hussain, A.; El-Tabakh, M.A. & Khan, S.A. 2024. Diversity and genetic structure of freshwater shark Wallago attu: An emerging species of commercial interest. Environmental Science and Pollution Research 31(10): 15571-15579.
- Sohail, M., Kamran, M.A.; Mansha, S.; Nawaz, A. & Bashir, F. 2025. Epidermal antimicrobial peptides in fish and frogs: from natural defense to biomedical applications. Journal of Wildlife and Ecology 9(3): 333-351.
- Strauss, R.E. & Bookstein, F.L. 1982. The truss: Body form reconstructions in morphometrics. Systematic Zoology 31(2): 113-135.
- Talwar, P.K. & Jhingran, A.G. 1991. Inland fishes of India and adjacent countries (Vols. 1–2). Oxford & IBH Publishing.
- Turan, C. 2004. Stock identification of Mediterranean horse mackerel (Trachurus mediterraneus) using morphometric and meristic characters. ICES Journal of Marine Science 61(5): 774-781.
- Wootton, R. J. (1990). Ecology of teleost fishes. Chapman & Hall.
- Zamani-Faradonbe, M.; Keivany, Y.; Abbasi-Jeshvaghani, M. & Asadi-Namavar, M. 2020. Morphometric and meristic variation in twelve different populations of Garra rufa (Heckel, 1843) from Iran. Jordan Journal of Natural History 7: 108-124.
- Zohra, K.; Osmany, H. & Imran, H. 2026. Seasonal Variation in Feeding Habit of Grey Sharpnose Shark (Rhizoprionodon oligolinx) Springer, 1964 from the Northern Arabian Sea. Journal of Wildlife and Ecology 9(4): 394-403.
References
Ali, A., Shah, S.; Ali, A. & Haris, M. 2024. Fish diversity in different streams of Lower Swat, Pakistan. Journal of Wildlife and Ecology 8: 144-153.
Altaf, M. 2026. Freshwater Fishes of Pakistan: Ecology, Distribution and Threats. Journal of Wildlife and Ecology. 10(SI-1).
Bagenal, T.B. & Tesch, F.W. 1978. Age and growth. In T. B. Bagenal (Ed.), Methods for assessment of fish production in fresh waters (3rd ed., pp. 101-136). Blackwell Scientific Publications.
Basharat, H.; Ghafoor, A.; Chavhan, A.; Zafar, M.; Abbas, K. & Parveen, J. 2016. Microsatellite markers revealed poor genetic structure of Wallago attu in Punjab, Pakistan. International Journal of Life Sciences 4(3): 385-393.
Begg, G.A. & Waldman, J.R. 1999. A holistic approach to fish stock identification. Fisheries Research 43(1-3): 35-44.
Benjamini, Y. & Hochberg, Y. 1995. Controlling the false discovery rate: A practical and powerful approach to multiple testing. Journal of the Royal Statistical Society: Series B (Methodological) 57(1): 289-300.
Cadrin, S.X. 2000. Advances in morphometric identification of fishery stocks. Reviews in Fish Biology and Fisheries 10(1): 91-112.
Franssen, N.R. 2011. Anthropogenic habitat alteration induces rapid morphological divergence in a native stream fish. Evolutionary Applications 4(6): 791-804.
Grill, G.; Lehner, B.; Thieme, M.; Geenen, B.; Tickner, D.; Antonelli, F.; Babu, S.; Borrelli, P.; Cheng, L.; Crochetiere, H. & Ehalt Macedo, H. 2019. Mapping the world's free-flowing rivers. Nature 569(7755): 215-221.
Haas, T.C.; Blum, M.J. & Heins, D.C. 2010. Morphological responses of a stream fish to water impoundment. Biology Letters 6(6): 803-806.
Habib, S.S.; Naz, S. & Mehmood, R. 2020. Morphometric relationships of some specified species of the family Cyprinidae in Jinnah Barrage, Punjab, Pakistan. Journal of Bioresource Management 7(2).
Hetzel, C. & Forsythe, P. 2023. Phenotypic plasticity of a generalist fish species resident to lotic environments: Insights from the Great Lakes region. Ecology and Evolution 13(11): e10715.
Imran, H., Zohra, K. & Osmany, H. 2025. Stomach analysis of the Milk shark, Rhizoprionodon acutus (Ruppell, 1837) in the Coastal Waters of the Pakistan, Northern Arabian Sea. Journal of Wildlife and Ecology 9: 268-281.
Kalsoom, B.; Mishaal, B.; Sardar, T.; Nasreen, Z.; Kalsoom, S. & Sharif, A. 2021. Water quality parameters and fish diversity at Chashma Barrage, Mianwali, Punjab, Pakistan. RADS Journal of Biological Research & Applied Sciences 12(2): 135-145.
Kelley, J.L.; Davies, P.M.; Collin, S.P. & Grierson, P.F. 2017. Morphological plasticity in a native freshwater fish from semiarid Australia in response to variable water flows. Ecology and Evolution 7(16): 6595-6605.
Klingenberg, C.P. 2022. Methods for studying allometry in geometric morphometrics: A comparison of performance. Evolutionary Ecology 36(4): 439-470.
Kumar, A.; Negi, N.; Yadav, N.; Badola, R.; Hussain, S.A. & Gupta, S.K. 2024. Genetic diversity, spatial connectivity, and population structure of Asian silurid catfish Wallago attu (Bloch and Schneider, 1801) in the Ganga River System: Insights from mitochondrial DNA analysis. Molecular Biology Reports 51(1): Article 378.
Kumar, A.; Negi, N.; Yadav, N.; Badola, R.; Hussain, S.A. & Gupta, S.K. 2024. Genetic diversity, spatial connectivity, and population structure of Asian silurid catfish Wallago attu (Bloch and Schneider, 1801) in the Ganga River System: Insights from mitochondrial DNA analysis. Molecular Biology Reports 51, Article 378.
Kumar, G.; Kashyap, A. & Serajuddin, M. 2023. Assessment of morphometric variations among the populations of Asian sheat catfish Wallago attu (Siluridae) from five Indian rivers. Inland Water Biology 16(6): 1026-1037.
Kwikiriza, G.; Yegon, M.J.; Byamugisha, N.; Beingana, A.; Atukwatse, F.; Barekye, A. & Meimberg, H. 2023. Morphometric variations of Nile tilapia (Oreochromis niloticus) (Linnaeus, 1758) local strains collected from different fish farms in South Western Highland Agro-Ecological Zone, Uganda: Screening strains for aquaculture. Fishes 8(4): 217.
Langerhans, R.B. 2008. Predictability of phenotypic differentiation across flow regimes in fishes. Integrative and Comparative Biology 48(6): 750-768.
Langerhans, R.B. & Reznick, D.N. 2010. Ecology and evolution of swimming performance in fishes: Predicting evolution with biomechanics. In P. Domenici & B. G. Kapoor (Eds.), Fish locomotion: An eco-ethological perspective (pp. 200-248). Science Publishers.
Le Cren, E.D. 1951. The length–weight relationship and seasonal cycle in gonad weight and condition in the perch (Perca fluviatilis). Journal of Animal Ecology 20(2): 201-219.
Malik, D.S.; Sharma, A.K.; Sharma, A.K.; Thakur, R. & Sharma, M. 2020. A review on impact of water pollution on freshwater fish species and their aquatic environment. In Advances in Environmental Pollution Management: Wastewater Impacts and Treatment Technologies 1: 10-28.
Mouludi-Saleh, A.; Keivany, Y. & Jalali, S.A.H. 2018. Comparison of meristic traits in Transcaucasian chub (Squalius turcicus De Filippi, 1865) from Caspian Sea basin. International Journal of Aquatic Biology 6(1): 8-14.
Mirza, M. & Sandhu, A. 2007. Fishes of the Punjab, Pakistan. Lahore, Pakistan: Polymer Publications.
Muhammad, N. & Zain-ul-Abideen, S.M. 2022. Diversity of fishes of head Marala, Punjab, Pakistan. Journal of Wildlife and Ecology 6(2): 58-63.
Muhammad, N.; Rasool, F. & Imran, M. 2026. Mitochondrial ATPase 6/8-Based Assessment of Genetic Diversity and Population Structure in Cirrhinus mrigala Across the Indus River System, Pakistan. Journal of Wildlife and Ecology 10(SI-1).
Muhammad, N.; Altaf, M.A.; Abbasi, M.; Khan, A.M. & Iqbal, K.J. 2023. From water to Remedy: Fishes as Ethnopharmacological Resources along the River Ravi, Punjab, Pakistan. Journal of Wildlife and Ecology 7(3): 93-108.
Muhammad, N.; Imran, M. & Abideen, S. 2024. Exploring the fish diversity along the River Ravi in Punjab, Pakistan. Journal of Wildlife and Ecology 8: 74-81.
Naseer, A.; Imran, M.; Khan, A.M. & Ammar, M. 2023. A comparative morphologic and craniometric analysis of grass carp (Ctenopharyngodon idella) population from Punjab, Pakistan. Journal of Xi’an Shiyou University 19(4): 48-62.
Ng, H.H.; de Alwis Goonatilake, S.; Fernado, M. & Kotagama, O. 2020. Wallago attu. The IUCN Red List of Threatened Species. International Union for Conservation of Nature.
Osmany, H.B.; Zohra, K. & Imran, H. 2026. Seasonal Variation in Feeding Habits of the Hardnose Shark (Carcharhinus macloti) and Comparison of Diet with Other Demersal Sharks in the Northern Arabian Sea of Pakistan. Journal of Wildlife and Ecology 10(SI-1).
R Core Team. 2023. R: A language and environment for statistical computing. R Foundation for Statistical Computing.
Rawat, S.; Benakappa, S.; Kumar, J.; Naik, A.S.K.; Pandey, G. & Pema, C.W. 2017. Identification of fish stocks based on truss morphometric: A review. Journal of Fisheries and Life Sciences 2(1): 9-14.
Reist, J. D. 1985. An empirical evaluation of several univariate methods that adjust for size variation in morphometric data. Canadian Journal of Zoology 63(6): 1429-1439.
Rohlf, F.J. 2015. The tps series of software. Hystrix, the Italian Journal of Mammalogy 26(1): 9-12.
Secer, B.; Mouludi-Saleh, A.; Eagderi, S.; Poorbagher, H.; Cicek, E. & Sungur, S. 2022. Phenotypic plasticity of angora loach, Oxynoemacheilus angorae (steindachner, 1897) in inland waters of turkey. Iranian Journal of Science and Technology, Transactions A: Science 46(5): 1317-1326.
Sajjad, A.; Jabeen, F.; Ali, M. & Zafar, S. 2023. DNA barcoding and phylogenetics of Wallago attu using mitochondrial COI gene from the River Indus. Journal of King Saud University–Science 35(6): 102725.
Shabir, M., Bano, N. & Majeed, U. 2025. Effect of Fermented Moringa (Moringa oleifera) Leaves as a Feed Ingredient in the Diet of Rohu (Labeo rohita) Fish. Journal of Wildlife and Ecology 9(1): 31-42.
Sherzada, S.; Hussain, N.; Hussain, A.; El-Tabakh, M.A. & Khan, S.A. 2024. Diversity and genetic structure of freshwater shark Wallago attu: An emerging species of commercial interest. Environmental Science and Pollution Research 31(10): 15571-15579.
Sohail, M., Kamran, M.A.; Mansha, S.; Nawaz, A. & Bashir, F. 2025. Epidermal antimicrobial peptides in fish and frogs: from natural defense to biomedical applications. Journal of Wildlife and Ecology 9(3): 333-351.
Strauss, R.E. & Bookstein, F.L. 1982. The truss: Body form reconstructions in morphometrics. Systematic Zoology 31(2): 113-135.
Talwar, P.K. & Jhingran, A.G. 1991. Inland fishes of India and adjacent countries (Vols. 1–2). Oxford & IBH Publishing.
Turan, C. 2004. Stock identification of Mediterranean horse mackerel (Trachurus mediterraneus) using morphometric and meristic characters. ICES Journal of Marine Science 61(5): 774-781.
Wootton, R. J. (1990). Ecology of teleost fishes. Chapman & Hall.
Zamani-Faradonbe, M.; Keivany, Y.; Abbasi-Jeshvaghani, M. & Asadi-Namavar, M. 2020. Morphometric and meristic variation in twelve different populations of Garra rufa (Heckel, 1843) from Iran. Jordan Journal of Natural History 7: 108-124.
Zohra, K.; Osmany, H. & Imran, H. 2026. Seasonal Variation in Feeding Habit of Grey Sharpnose Shark (Rhizoprionodon oligolinx) Springer, 1964 from the Northern Arabian Sea. Journal of Wildlife and Ecology 9(4): 394-403.
