The present study investigated alterations in the structure, diversity and richness of fish species communities during three different seasons, while also comparing physico-chemical characteristics of the Dhansiri River, one of the largest tributaries of Brahmaputra. Spatio-temporal changes in environmental factors are key determinants of freshwater fish assemblages, influencing habitat structure and species composition (Fischer and Paukert 2008; Elías et al. 2020). This study offers valuable insights into ecological shifts within the river, enhancing the understanding of the intricate interrelationships between fish populations and their habitat. The Dhansiri River is characterized by high DO levels, high total alkalinity and an alkaline pH, which are comparable to other hilly rivers in the region (Gurumayum et al. 2014; Sarmah et al. 2020; Lkr et al. 2020). Spatial analysis of water quality parameters revealed that the upper stretch (S1) reflects a relatively pristine habitat with minimal human impact. On the contrary, a slight deterioration in water quality was noticed downstream, with comparatively poorest condition at S4 near Dimapur town. This stretch of the river is characterized by low DO levels, high CO2 and presence of NH3, NO2 and NO3. Stretch of the Dhansiri near Dimapur town has been listed as polluted and placed under Priority I category due to high BOD load (Anon 2019). Poor downstream environment quality may be attributed to increased human habitation, high anthropogenic discharge, agricultural runoff and changes in land use patterns (Premke et al. 2020; Ji et al. 2021). Pristine nature of water at S1 may be explained by its location within INP, where human interference is restricted. Distinct seasonal changes in current patterns, discharge rates, and temperature bring about unique alterations in the physico-chemical properties of riverine ecosystems (Pradhan et al. 2009). These seasonal changes affect the distribution of nutrients and dissolved gases, which in turn influence biological productivity and the species composition of the river.
PCA analysis indicated the importance of water chemistry, such as pH and total alkalinity in determining primary axis of water quality variation (Saalidong et al. 2022). The dominance of these parameters suggests that river’s buffering capacity and its ability to neutralize acidic inputs are critical to maintaining its ecological balance. Additionally, the significant presence of NO2, NO3 and TDS as influential contributors highlights the role of nitrogen compounds (Vorobyeva et al. 2021) and dissolved solids in shaping water quality profile and its biological productivity. Excess nitrogen in aquatic ecosystems can lead to eutrophication, which disrupts aquatic food webs and reduce biodiversity (Dodds and Smith 2016). Furthermore, water temperature also emerged as a critical abiotic factor influencing Dim2. Relatively smaller contributions of other variables to Dim2 indicate their secondary importance in explaining water quality variation.
The study revealed a total of 69 fish species, distributed across 20 families and 46 genera, indicating significant diversity of freshwater fishes comparable to other major river systems in India (Sarkar et al. 2010; Shukla and Bhat 2017). Highest number of species was recorded along the upper stretch of river at S1 (54), while the least was observed along the lower stretch at S4 (37). The upper stretch (S1) is located within the Intangki National Park, an authorized biodiversity conservation area. This site is restricted from anthropogenic interference and features a pristine ecosystem with high diversity, contributing to the observed richness and diversity of fish species. Conversely, the lower stretch (S4) near Dimapur town showed significant declines in fish diversity. This decline can be attributed to the deterioration of environmental variables, as recorded in the study (Table 3), along with the decline in structural habitat complexity, and the increased human habitation and activities observed during downstream sampling. Poor environmental parameters (lower DO, higher NH₃ and higher CO₂) (Anon 2019) along with increased anthropogenic pressure (e.g. pollution, over exploitation etc.) may have negatively impacted fish species richness, diversity, distribution and relative abundance. These findings are consistent with previous studies showing that degraded water quality and habitat deterioration adversely affect freshwater fish populations (Anon 2019; Dudgeon et al. 2006). Typically, in riverine systems species richness, diversity, and abundance increase from upper to lower stretches due to broad range of habitats and resources available (Weber and Peter 2007). However, the Dhansiri River exhibited an opposite trend, with species numbers declining from upstream to downstream. The observed spatial variation in diversity, number, and distribution of fish species is attributed to several factors, including natural topography, environmental influencers (Hashemi et al. 2015), pollution (Anon 2019), habitat complexity (Smokorowski and Pratt 2007), reduced water quality and increased anthropogenic pressures, which limit the suitability of downstream habitats for diverse fish communities (Table. 3) (Lakra et al. 2010; Pandey and Radhakrishnan 2022).
Seasonally, a higher number of fish species was observed in the Dhansiri River during monsoon at all stations, with the exception of S2 (pre-monsoon). Water discharge has been identified as the single most important factor driving fish species richness in Himalayan rivers (Bhat et al. 2012). Increased water discharge during monsoon creates new habitats and improves connectivity between different habitats and sections of the river, facilitating fish migration and colonization of new areas, thereby increasing likelihood of encountering higher number of species. Higher number of fish species observed at S2 during pre-monsoon may be attributed to increased natural food availability (Acharjee and Barat 2014). Previous studies across rivers in south and Southeast Asia have consistently reported cyprinids as most dominant group of fishes (Bhat 2003). The results of this study, which observed that cyprinids exhibit the highest relative abundance compared to other groups is a testimony to this statement. Cyprinids dominate fish community structure in tropical Indian rivers owing to their high adaptive variability and ability to occupy heterogeneous habitats (Johnson and Arunachalam 2009).
Diversity indices in general are functions of different species available in an ecosystem and relative abundance of each species. Shannon diversity index is widely used to quantify diversity or richness of species in a specific ecological community and is one of the most commonly employed indices for comparing species diversity across habitats (Clarke and Warwick 2001). Margalef richness index is a biodiversity metric used to assess species richness in an ecological community (Margalef 1958). Simpson dominance index assesses the dominance or evenness of species abundances within an ecological community (Simpson 1949). On a spatial scale, values of ‘H′’ and ‘d’ were highest in pristine upper stretch (S1) with a gradual decline downstream. Conversely, values of ‘D’ increased from upstream to downstream, indicating a higher dominance of fewer species. Highest values of ‘H′’ (3.750) and ‘d’ (9.875) and lowest value of ‘D’ were observed during monsoon at S1, highlighting a high level of species richness and uneven distribution. This aligns with established ecological principles, as fish typically migrate, spawn and recruit during monsoon, leading to increased diversity and abundance in the ecosystem (Sreekanth et al. 2016). Restricted human interference by virtue of location and congenial habitat might be major factors for migration and breeding of fish during monsoon, leading to higher fish diversity and low dominance at S1. Pielou's evenness index quantifies the distribution of species within a community, providing a measure of how evenly individuals are spread across species. The index ranges from 0 to 1, where 0 represents maximum species dominance (only one species present), and 1 represents maximum evenness (all species are equally abundant) (Pielou 1966). In this study, it was observed that upper stretch (S1) has an even distribution of fish species, followed by S4. It should be noted that instances of high abundance in certain species may result in a deviation from evenness in overall species diversity (Saha et al. 2022), which might be one of the possible reasons for lower evenness in S2 and S3.
Cumulative dominance curve (k-dominance), which ranks fish species based on their abundance and the proportion of individuals per species, plotted against the logarithmic rank of species (Clarke 1990). The curved showed S4 as least diverse site, with a steep dominance curve indicating a few species dominating the community. Concept of species richness is commonly utilized as an indicator of environmental quality, as it tends to rise in presence of healthy ecosystems and minimal anthropogenic impairment (Roth et al. 2000; Kirk and Rahel 2022). Conversely, a decrease in species richness accompanied by proliferation of a few dominant species indicates a progressively deteriorating environment (Jacinto et al. 2023). It is important to note that freshwater assemblages are not solely affected by species zonation or spatial arrangement, but also shaped by physical disturbances, and habitat modifications (Daufresne and Boet 2007; Perkin et al. 2015). NMDS is an ordination technique used to visualize the similarity between samples based on ecological data. In the present study NMDS analysis revealed a clear separation of S4 from the other three stations. As per guidelines forwarded by Clarke (1993) stress values < 0.05 are considered excellent; < 0.10, good; < 0.20, useable; and > 0.20, not reliable. Lower the stress value, better the fitting in ordination plot. In the present study, a stress value of 0.07 indicates a good fit for the distances/dissimilarities in the spatial ordination plot.
Environmental factors play a crucial role in maintaining fundamental habitat characteristics and ecosystem functions, and they are known to significantly influence fish community structure in aquatic systems (Mondal and Bhat 2020; Debnath et al. 2022). Understanding the impact of environmental influencers on fish communities is essential for elucidating fish-habitat relationships, which can contribute to more effective ecosystem management and preservation (Gillooly et al. 2001; Wang and Lyons 2003; Li et al. 2018). BIO-ENV analysis found strong correlations between environmental parameters, fish abundance and composition, with NH3, NO3, CO2, TDS, TA, pH, specific conductivity, DO and WT being the most influential factors. Highest degree of correlation with fish abundance was observed in the combinations of NO3, CO2 and TDS. Environmental variables affects fish abundance and composition both directly or indirectly by influencing dynamics of fish food organisms such as plankton (Gogoi et al. 2019) and aquatic invertebrates (Das et al. 2022). WT is one of the critical influencers that affect distribution of fishes, both directly by influencing their metabolic and physiological processes (Gillooly et al. 2001) and indirectly by affecting biological productivity and food matter availability (Sarkar et al. 2021). Additionally, TDS, TA and pH have been shown to reportedly influence chlorophyll contents in rivers (Sarkar et al. 2021), which are closely associated with productivity and the availability of food sources, thereby affects fish distribution. Similarly, NH3, NO3, CO2 are also associated with biological productivity of aquatic ecosystems (Galloway et al. 1994; Sigman and Hain 2012) and can directly influence fish community structure. Parameters like pH, WT, TDS, NO3, DO are known to influence dynamics of fish food organisms (Gogoi et al. 2019; Das e al. 2022), which in turn affects distribution of fishes. Furthermore, DO, pH, NH3, NO3, CO2, TA have direct effects on fish metabolism, which may influence their distribution and abundance (Boyd and Tucker 1998). It is to be noted that, DO is the single most critical parameter for the survival of all forms of aquatic life (Keke et al. 2016) and it therefore exerts influence on fish communities. Managing adequate DO levels is essential for sustaining healthy fish populations and preserving the overall balance of aquatic ecosystems.
The present study highlighted the impact of human activities, particularly at S4 near Dimapur town, as evidenced by reduced species richness, higher dominance of fewer species, and poorer water quality parameters. The presence of higher concentration of NH₃, NO₃ and CO₂ at S4 reflects inputs from agricultural runoff, urban wastewater and industrial discharges, all of which contribute to the degradation of Dhansiri River ecosystem. The finding of this study underscore the need for implementation of effective regulation and management measures to protect this Dhansiri River’s biodiversity, particularly in downstream areas. Establishing buffer zones, improving wastewater treatment and controlling agricultural runoff are critical steps towards mitigating the negative impact of anthropogenic activities (Hughes et al. 2018). Additionally, the presence of several species listed as Near Threatened and Vulnerable highlights the importance of targeted conservation efforts. Protecting critical habitats, especially in the upper stretches of the river, is essential for maintaining the ecological integrity of the entire system. Further research on the population dynamics of these threatened species is necessary to inform conservation strategies and ensure their long-term survival. As previously mentioned, environmental aspects participate crucially in shaping fish communities. Given the current state of rapid urbanization and climate change, it is imperative that we gain critical insights into ecological mechanisms affecting fish communities in order to develop effective protection and conservation strategies. The present study, conducted in a least-explored tropical river from a globally significant biodiversity hotspot, contributes to our knowledge and understanding, providing cues for sustainable riverine fisheries management.