River Ecosystems of the Central Highland Ecoregion: Spatial

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River ecosystems of the Central Highland ecoregion: Spatial distribution of benthic ﬂora and fauna in the Plateau rivers (tributaries of the Yamuna and Ganga) in Central India Prakash Nautiyal , 1,* Asheesh Shivam Mishra,1 Jyoti Verma,2 and Asha Agrawal3 1Aquatic Biodiversity Unit, Department of Zoology and Biotechnology, HNB Garhwal University, Srinagar 246174, Uttarakhand, India 2Department of Zoology, University of Allahabad, Allahabad 211002, Uttar Pradesh, India 3Department of Zoology, Sri Sathya Sai College for Women, Bhopal, Madhya Pradesh, India *Corresponding author: [email protected]

This study reveals the relationship of biotic assemblages to their environments in the Central Highland Ecoregion rivers (Ken, Paisuni and Tons). Such knowledge will enhance our predictive abilities in the ecological studies and resource management in the concerned region. Rivers of an ecoregion are expected to harbour similar ﬂora, fauna, and communities. This hypothesis was examined by tracking the spatial variation in richness, density, and taxonomic composition of benthic diatom and macroinvertebrate assemblages in three Plateau rivers whose course lay within 3N latitude. Taxonomic richness and density decreased with the reduction of substrate size from the headwaters to the lowlands. Cluster and ordination analyses indicate prevalence of intrabasin rather than interbasin gradients of abundance. In both assemblages, the most abundant taxon is rarely similar longitudinally or across the river sections in the ecoregion. Hence, proximate factors govern the assemblages rather than ecoregional properties (latitude, altitude). This, and the high richness of diatom ﬂora, points to diverse assemblages from the headwater to the lowland section in each river and among the rivers of the ecoregion.

Keywords: benthic assemblages, richness, ordination, river-link, trophic state

Introduction linkages. Ecoregions may poorly classify compo- nents of the biota that are strongly regulated by local Correspondence between ecoregional classiﬁca- factors. Algae, macroinvertebrate and ﬁsh assemb- tion and the biota should depend on the strength of lages each respond to environmental changes at

Color versions of one or more of the ﬁgures in the article can be found online at www.tandfonline.com/uaem.

Aquatic Ecosystem Health & Management, 20(1–2):43–58, 2017. Copyright Ó 2017 AEHMS. ISSN: 1463-4988 print / 1539-4077 online DOI: 10.1080/14634988.2017.1296324

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different temporal and spatial scales because of their communities of the Ken, Paisuni, and Tons Rivers different life histories, physiologies, and mobilities. during the last decade help predict ecoregional fea- Thus, changes in each assemblage may reflect local tures of the Central Highlands (Nautiyal and Mishra, conditions of stream habitats including microhabi- 2012, 2013; Mishra and Nautiyal, 2013a; 2013b). tats, channel unit, reach, and watersheds (Frissel The benthic diatoms and macroinvertebrates et al., 1986). Diatom assemblages often respond to (usually insect larvae) account for the majority of land use, especially agricultural activities (Pan species among the producers and consumers. A et al., 2000). Thus, diatom assemblages not specific profitable fishery depends on healthy food webs to bioregions may be ideal as unbiased indicators of that involve lower trophic levels within benthic stream water quality (Charles et al., 2006). The dis- communities. Fishing is a commercial activity that tribution of benthic macroinvertebrates also varies generates considerable employment along the riv- within and among the rivers (Hawkins and Sedell, ers, from those manufacturing fishing nets, to fish- 1981; Milesi et al., 2009; Mesa, 2010; Ezekiel ing vessels, fishermen, and traders. This study et al., 2011). Macroinvertebrate compositions are primarily examines the richness and composition known to exhibit much stronger relationships to of benthic diatom and macroinvertebrate commu- local environmental conditions (at reach scale) than nities and the factors governing them in the Cen- to catchment variables (Carter et al., 1996). The fac- tral Highlands ecoregion. The focus of this study tors influencing freshwater biodiversity at local and was to generate information on the spatial distribu- regional scales in India (Rodgers et al., 2002) tion of benthic biota along the length of the rivers. remains poorly understood. Our study area is within the Central Highlands subdivision of the major physiographic region the Materials and methods Peninsular Plateau, synonymous with the 6th biogeographic region—Deccan Peninsula. The Central Study area Highlands alongside Chota Nagpur are the north- ernmost biotic provinces of this biogeographic region The Ken, Paisuni, and Tons drainages in the (Source: wiienvis.nic.in/database/htmlpages/biopro- northeastern Central Highlands are the subject of vincemap.htm WII, 2000). The Central Highlands this study. The Central Highlands consist of the province has an east–west orientation and continental Malwa, Bundelkhand and Chota Nagpur Plateaus climate, with rainfall decreasing westwards. The that form the northern subdivision of the ancient Gangetic drainage primarily descends from the (Gondwana) triangular-shaped tableland—the Himalayas as the Yamuna and Ganga Rivers. The Peninsular Plateau. This geographical region is Plateau Rivers, descending from the western and cen- bound by the Gangetic Plains to the north and east, tral Vindhya Ranges, join the Yamuna, while those and the Deccan Peninisula to the south. The Vind- arising from the eastern ranges join the Ganga. The hya Ranges give rise to northward flowing Central Highlands is an important region for the tributaries of the Yamuna (Chambal, Kali, Sindh, freshwater biodiversity of hill streams of the Indian Parbati, Betwa, Dhasan, Ken, Paisuni) and Ganga subcontinent. This ecoregion represents a vital junc- (Tons, Sone) Rivers. These rivers are important tion of mountain streams and rivers. for irrigation, domestic supply, and power genera- Agenda 21 (1992) emphasizes the role of biodi- tion including proposed river linking projects on versity in the sustainable development of the uplands. the Indian Peninsula (NWDA, 2006). The rivers Only recently has biodiversity been documented for also have high religious significance. These drain- the Central Highlands ecoregion (Nautiyal and ages lie within 24 to 26N latitude, 79 to 82E lon- Verma, 2009a; Verma and Nautiyal, 2010; Nautiyal gitude, and altitude 360 to 72 m above sea level and Mishra, 2012, 2013; Kulshrestha, 2009; Lakra from source to confluence. The examined length et al., 2010; Dwivedi and Nautiyal, 2012; Nautiyal of rivers Ken, Paisuni, and Tons are ca. 340 km, et al., 2011). Stable native food webs are the key to 100 km, and 305 km long, respectively. Locations health of the ecosystem. Lower trophic levels in wild sampled on the Ken are labelled K1 to K4, Paisuni rivers, especially grazers, require pristine conditions. P1 to P4, and Tons T1 to T4 (Figure 1), dividing It is imperative to document the baseline conditions the river into four gross sections: upper (K1, P1, of benthic biota and assemblages (diatom, macroin- T1), middle (K2, P2, T2), and lower plateau (K3, vertebrates). Recent studies of the benthic P3, T3), and a lowland section (K4, P4, T4). The

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Figure 1. Central highlands drainage. The rectangular marked area is the location of study area in the Gangetic basin. The map on left top displays terrain and main course of the Central Highlands rivers, primarily the Betwa, Ken, Paisuni and Tons. The terrain map depicts Vindhyan ranges as dark broken patches from where rivers and streams descend ﬁrst into the Plateau and then the Gangetic Plains. The large box in the centre of the terrain map encompasses the location of the rivers Ken, Paisuni, and Tons and the smaller box to left displays the course of the Betwa River in respect to the Ken. The map on right shows the sampling stations selected to represent various sections: Headwater section (K1, P1, T1), M: Middle section (K2, P2, T2), L: lower section (K3, P3, T3), Mo: Lowland section (K4, P4, T4). Acronyms: K: Ken, P: Paisuni, T: Tons. The map (left below) depicts the position of the Betwa in respect to the Ken and location of the proposed Ken–Betwa link.

course of these rivers from upper to lower sections June) was considered appropriate. The river’s flora moves through the Plateau, which constitutes the and fauna diversify after monsoon floods (July to highland sections, while the mouths fall in the September). The dry period extends from October to lowland section, which stretches for approxi- June but sampling was restricted to December 2003 mately 80 km in the Ken and Tons, and 30 km in to March 2004 because long stretches of the rivers the Paisuni. The highland stretch of the Ken and tendtodryupduringthesummerseason(Marchto Tons Rivers consists of short headwater sections June), forming pools of various sizes that disrupt the up to 50 km, and 100–120 km long middle and continuum. lower sections. In contrast, the Paisuni headwater Water temperature and pH were analyzed with section is approximately 10 km and the middle portable meters (WT-Mextech, multimeter, pH- and lower sections are 30–40 km. The physiogra- Hanna). Current velocity (CV) was measured by phy, climate, and vegetation have been described an EMCON current meter. Substrate predomi- previously for these rivers (Mishra and Nautiyal, nance at each sampling station was estimated visu- 2011; Nautiyal and Mishra, 2012). ally. If the river bed was extensively littered with cobble substrate followed by boulder and gravel, then substrate type was recorded as C;B:G. Sub- Sampling strate names and their categories were determined through Wentworth classification of substratum Annual intensive sampling is considered suitable particle size (http://fwcb.cfans.umn.edu/courses/ to analyze the distributional patterns of benthic mac- fw8459/Private/SubstrateandHyporheos08-2.pdf). roinvertebrate assemblages (Corkum, 1989). Thus, Diatom samples were obtained by scraping the intensive sampling in the dry period (October to surface of 3£3 cm samples of cobbles collected

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from the river bed Samples were collected from 7.0–7.8). The range value of current velocity (CV) various habitats at each station. Each sample was was higher in the Ken (1.0–60 cms¡1) and Tons preserved in 4% formaldehyde solution. Samples (0.1–50 cms¡1) compared with the Paisuni (0.0– were treated with HCl and cleaned by boiling in 30.9 cms¡1). The WT and pH increase down- H2O2. The treated samples were washed repeat- stream of the source, while current velocity edly to remove traces of acid and peroxide. Perma- decreases (Figure 2). Substrate particle size nent Naphrax mounts were prepared and examined decreases gradually from the upper to lower pla- under bright field using a BX-40 Trinocular Olym- teau sections, but abruptly changes to small sedi- pus microscope (x10 and x15 wide-field eyepiece) ments in the lowland section, which lies in the fitted with a PLANAPO x100 oil immersion objec- Gangetic Plains (Table 1). tive. Diatoms were identified according to Hustedt (1931–1959), Krammer and Bertalot (1986–1991), Sarode and Kamat (1984), and Gandhi (1998). Benthic diatoms The slides are archived at the Aquatic Biodiversity Unit. The relative abundances (in percent) were The diatom flora is represented by 205, 202 and determined on the basis of 300 valves from each 211, taxa, respectively, in the rivers Ken, Paisuni and sample at each station. Tons, with a total overall 293 taxa (species, variety, Benthic macroinvertebrate fauna was sampled form) from 50 genera (see Appendix A in the online intensively (20 quadrants per station). Sampling pro- cedures at each station involved lifting stones (boulder, cobble, pebble, gravel) and sieving clay and silt from a 0.09 m2 area in different flows (turbulent, swift, slow, placid). The substrate was washed to dis- lodge the fauna, which was preserved in 5% formalin for further analysis. As broad taxonomic classifica- tions are acceptable (Corkum, 1989), the fauna was identified at a family level (Edmondson, 1959; Eding- ton and Hildrew, 1995). Counts were made to obtain density (indiv. m¡2) and relative abundance. The functional groups were defined according to Cotta Ramusino et al. (1995) and Dudgeon (1984). The Kruskal–Wallis test (Henderson, 2003) was applied to determine significant differences between stations in total density of the community and higher taxa (order, families). Principal Component Analysis (PCA) determined the characteristic taxa at each station, and Canonical Correspondence Analysis (CCA; ter Braak and Smilauer, 2002) helped to identify the environmental variable(s) causing longitudinal variation in the taxonomic composition of the benthic macroinvertebrate fauna. The cluster analysis to classify river sites and the multivariate analysis (CCA and PCA) were based on the raw data, i.e. counts of diatom species and invertebrate fauna from each quadrant.

Results The range of water temperature (WT) and pH differs in the rivers Ken (16–27 C; 7.0–7.5), Pai- Figure 2. Longitudinal proﬁle of physicochemical characteris- suni (15.5–33 C; 7.0–7.7) and Tons (17–31.4 C; tics in the rivers Ken, Paisuni and Tons.

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Table 1. Physiographic features of various sampling stations in the Vindhyan rivers: Ken, Paisuni and Tons.

River/ Altitude Distance from Land Station (m asl) Source (km) Use Substratum Ken River Shahnagar (K1) 365 ca.10 Agriculture- Rock; Boulder; Cobble; Pebble Village Panna (K2) 200 142.5 Forest- Boulder; Rock; Cobble; Pebble; Gravel Agriculture Banda (K3) 95 267.5 Agriculture Cobble; Pebble; Gravel; Boulder; Sand; -City Silt Chilla (K4) 86 340 Agriculture Clay-Silt Paisuni River Anusuya (P1) 180 10 Forest Rock; Boulder; Cobble; Pebble Chitrakut (P2) 135 26 Agriculture Clay; Silt; Boulder; Pebble-Gravel -Temple Purwa (P3) 131 42 Agriculture Silt; Cobble; Cobble-Pebble; Boulder; Sand; Pebble-Gravel; Clay Rajapur (P4) 80 100 Agriculture Silt; Cobble-Pebble; Clay; Sand; Pebble- Gravel Tons River Maihar (T1) 326 ca.56 Agriculture Boulder; Cobble; Pebble; Gravel; Silt Satna (T2) 290 98 Agriculture Cobble; Pebble; Pebble-Gravel; Sand; -Village Silt Chakghat (T3) 94 232.5 Agriculture Rock; Boulder-Pebble; Cobble; Pebble- -Town Gravel; Silt Meja (T4) 72 305 Agriculture Clay-Silt

supplemental material). In the Ken, richness declines In the ecoregion, 18 diatom taxa attain 10% or downstream of the headwater section, while in the more relative abundance in one or more sections. case of the Paisuni and Tons, richness increases from Amphora twentiana (9%) is also listed, as it is the headwater to middle sections but decreases from probably unique to the rivers of Central India. The the lower to lowland sections (Figure 3). Total den- middle section of the Tons contains eight taxa that sity decreases downstream of the sources in the Ken attained >10% abundance, compared with four and Tons, while in the Paisuni density increases from and two taxa, respectively, in the lower and low- the headwater to middle section and declines in the land sections (Table 2). Notably, all Achnanthi- lowland section (Figure 3). Richness as well as den- dium species occur in 9–14% abundance at all the sity vary among the headwater (168–182; 1198– locations of these rivers. Another 11 taxa exhibit 1404 cell. mm¡1), middle (178–185; 1098– >10% abundance at either the headwater or low- 1343 cell. mm¡1), lower (157–169; 803–1003 cell. land location/section. Aulacoseira granulata, mm¡1), and lowland (151–156; 843–934 cell. occurs in >10% relative abundance in all the sec- ¡ mm 1) sections of these rivers, most noticeably in tions of the Ken only. Based on abundance pat- the lowland sections compared with the headwater to terns, Achnanthidium chitrakootense was the lower sections. The raphid elements, especially bira- representative taxa for Paisuni River, while Ach- phid, constitute a very large share of diatom ﬂora nanthidium minutissimum var. scotica and A. min- throughout the ecoregion and in each basin. The bira- utissimum were representative taxa for the Ken phid Navicula, Cymbella,andNitzschia are species- and Tons, respectively. These taxa are, therefore, rich genera in each river (see Appendix B in the the abundance signatures of the respective rivers online supplemental material). (Table 2).

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Figure 3. Comparative view of longitudinal changes in species richness and density of benthic diatoms (primary vertical axis) and macroinvertebrates (secondary vertical axis).

Examination of the abundance pattern among the Paisuni (P1, P2). Similarly, the middle to lowland sections of the Ken, Paisuni, and Tons reveals the sections of the Ken (K2 to K4) resemble the lower similarity between the headwater section of the Ken section of the Paisuni (P3). The Tons forms a separate (K1), and headwater as well as middle section of the cluster of its own sections (Figure 4).

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Table 2. Distribution of diatom and macroinvertebrate (MI) abundance in the Plateau rivers. Diatom taxa with >10% and MI with highest percentage in any section of a river have been recorded as abundant irrespective of their abundance in other rivers or sections. The lowland zone of the Paisuni was not sampled. Acronyms: K, P, T D Ken, Paisuni, Tons; K1, P1, T1 D Headwater; K2, P2, T2 D Middle section; K3, P3, T3 D Lower section; K4, P4, T4 D Lowland section.

Benthic macroinvertebrates varies among the headwater, middle, and lower sections of these rivers. Richness is high in the Twenty-eight invertebrate taxa (class/fami- middle and lower sections of the Ken, the mid- lies), including those from seven insect classes, dle and lowland sections of the Paisuni, and the occur at the ecoregion scale, and 21, 24 and 27 headwater and lower sections of the Tons taxa are seen at the basin scale in the Ken, (Figure 3). Longitudinally, the mean density Paisuni and Tons, respectively. The richness decreased from the headwater to the lowland

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Figure 4. Similarity dendrogram (Bray–Curtis) based on diatom abundance counts for each taxon in each section of the rivers Ken, Paisuni and Tons. Acronyms as in Figure 1.

section in the Ken and Tons, while it increased the collectors (suspension and deposit feeders in the Paisuni (Figure 3). The density differed that filter fine particles from the water column or significantly among the sections (Observed gather loose particles in depositional areas) are H(0.5,) for headwater, middle, lower and lowland dominant longitudinally throughout the rivers, sections was 16.49, 13.76, 39.88 and 41.77, from the headwaters to the lowland sections. respectively). The percentage of scrapers (feeders of photosyn- Insect larvae are the major component of the thetic algae growing on any substrate) increases macroinvertebrate fauna followed by molluscs substantially downstream in the Ken and Tons, and annelids (Table 2). The order Ephemerop- while it decreases in the Paisuni (Figure 6). tera is a family-rich taxon in all of the Plateau rivers (Appendix C). The cluster analysis revealed that the invertebrate fauna reflect the Multivariate analysis close similarity among the middle, lower, and lowland (P2, P3, P4) sections of the Paisuni, as PCA ordination for diatoms reveals that well as the headwater, middle, and lowland Axes 1 and 2 represent 41.4% and 29.1% of the (T1, T2, T4) sections of the Tons. Only the low- variance in the lowland sections of the Ken and land sections of the Ken and Tons (K4-T4) are Tons (K4, T4). A. minutissimum, Gomphonema similar (Figure 5). The headwater and middle parvulum, Navicula viridula and Navicula cryp- sections of the Ken (K1 and K2) exhibit similar- totenella are characteristic taxa in these sec- ity with the lower section of the Tons (T3). Sim- tions. A. chitrakootense, A. petersenii, Navicula ilarly, the headwater section of the Paisuni (P1) seminulum and Navicula rostellata are charac- resembles the headwater and middle sections of teristic in the entire Paisuni and middle section the Tons (T1 and T2). The lower and lowland of the Tons (Figure 7). PCA ordination for sections of the Paisuni (P3 and P4) are similar to macroinvertebrate fauna indicates that Axes 1 the lower section of the Ken (K3). Functionally, and 2 represent 36% and 17.6% of the variance

Figure 5. Similarity dendrogram (Bray-Curtis) based on benthic macroinvertebrate fauna for the stations in the three rivers. Acro- nyms as in Figure 1.

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CCA indicated that land use, current velocity, and substratum are important variables for the distribution of benthic macroinvertebrate fauna in the rivers Ken, Paisuni, and Tons, respectively. However, depth (53.8%) and water temperature (46.2%) emerge as the most important factors with respect to the ecoregion (Figure 9).

Figure 6. Functional feeding groups in the Central Highlands rivers Ken, Paisuni and Tons. Acronyms: GC-Gathering collec- Discussion tor, FC-Filtering Collectors, P-Predator, SC-Scraper and Sh- Shredders. Benthic diatoms The richness of the diatom ﬂora for the ecore- in taxonomic composition. The characteristic gion is considerably higher than that found in a taxa are similar in the lowland sections of the lesser Himalayan stream (Verma and Nautiyal, Ken and Tons. The headwater of the Paisuni 2009) and the Plateau River Damodar (Nautiyal resembles the middle section of the Ken and Nautiyal, 1999a), but is comparable with what (Tabanidae, Sialidae) and faintly resembles the is seen in the Mandakini basin (Nautiyal et al., headwaters of the Ken and middle Tons, char- 2004a) and the Alaknanda-Ganga (Nautiyal and acterised by Glossoscolecidae, crustacean and Nautiyal, 1999b). When the 153 diatom taxa (38 Caenidae. The remaining sections of the Paisuni genera) recorded from the Betwa River (Figure 1, slightly resemble the lower section of the Ken, Appendix D) are added, the total number of taxa as Chironomidae lies intermediate to them. The in the Central Highlands increases to 322 repre- rivers Ken and Tons are similar compared with senting 50 genera (Appendix A). Kawecka and the Paisuni, with the exception of PI (Figure 8). Olech (2004) recorded 270 taxa from Finnish

Figure 7. PCA biplot to identify the characteristic taxa at each station of the Vindhyan rivers: Ken, Paisuni and Tons. Acronyms: A lin: A. linearis, Am: A. minutissimum,Ac:A. chitrakootense, A. petersenii, C div: C. diversa, C ex: C. excisa,Ctur: C. turgidula,G par: G. parvulum, A gr: A. granulata, N cry: N. cryptotenella, N rost: N. rostellata, N sem: N. seminulum, N vir: N. viridula and S u: S. ulna.

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Figure 8. PCA biplot to identify the characteristic taxa at the each station of the Vindhyan rivers. Acronyms: AG – Agrionida, BT – Baetidae, BR – Brachycentridae, CH – Chironomidae, CN – Caenidae, COR – Corbiculidae, DY – Dytiscidae, EP – Ephemer- illidae, GM – Gomphidae, GL – Glossosomatidae, GLO – Glossocolecidae, HE – Heleidae, HP – Heptageniidae, HL – Hydroptili- dae, HY – Hydropsychiidae, LP – Leptophlebiidae, NE – Neoephemeridae, NEP – Nepthydae, PR – Perlidae, RY – Rhyacophilidae, SA – Saliﬁdae, TB – Tabanidae and TH – Thiaridae.

Lapland, which was lower than the numbers seen Paisuni in proportion to its length supports the in the Central Highland ecoregion. Thus, the rich- view that freshwater biodiversity can be highly ness of the Central Highland rivers is higher than localized, since freshwater habitats are relatively that in the rivers of more temperate regions: 165 discontinuous and many freshwater species do not taxa found in the Estonian River (Vilbaste, 2001) disperse easily across the land barriers (WRI, 72 taxa in streams of Coastal Oregon (Naymik 1992). Nautiyal et al., (2004a) also observed ﬁve et al., 2005), and 98 and 94 taxa, respectively, centers of high diversity in different altitudinal from the Bulgarian rivers Vit and Osum (Stan- zones of the Mandakini basin. cheva et al., 2007). Species richness of the diatom community Richness is relatively low in the Paisuni and exhibits a general decrease from the headwater to Ken compared to the Tons. However, the shorter the lowland sections in all of the rivers, despite a Paisuni is proportionately richer than the longer slight increase in the middle sections of the Tons Ken and Tons Rivers. The island biogeography and Paisuni, attributed to the longitudinal gra- concept suggests that the larger area of the longer dients of decreasing substrate size and heterogene- rivers should have higher richness than the Pai- ity from source to mouth (Nautiyal and Verma, suni, provided the number of habitats increase 2009a). The richness of diatom ﬂora and density with area (Begon et al., 1990). However, diatom in the headwater and lower sections decreases richness did not increase with area, probably due towards the east from the Ken to the Tons basin, to the low richness in the lowland sections com- but increases in the middle and lowland sections pared with the upper plateau. High richness of the from the Ken to the Tons. The diatom richness in

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Figure 9. Canonical Correspondence Analysis (CCA) indicates the relationship between benthic macroinvertebrate taxa and environmental variables in the rivers Ken, Paisuni and Tons. The effective environmental variables are indicated in bold letters and the rivers in the ﬁlled triangle. Acronyms: ALT – Altitude CV – Current velocity, D – Depth, Dis – Discharge, DS – Distance from source, LAT – Latitude, LU – Land use, RW – River width, SUB – Substrate and WT – Water temperature. Other acronyms as in Figure 8.

the Paisuni is higher than in the river basins on Himalayan rivers, biraphid content is also high (Nau- either side, while density is slightly less. tiyal and Nautiyal, 1999b; Nautiyal et al., 2004b). The Central Highlands ecoregion supports high The Peninsular area or other parts of India, including Shannon species diversity (H) and low evenness: the mountainous zones like the Western Ghats Ken (5.01; 0.30), Tons (5.02; 0.33), and Paisuni (Krishnamurthy, 1954), are equally rich in biraphid (5.07; 0.23; Nautiyal and Verma, 2009b). The flora. Most species-rich genera are biraphid; Navi- Shannon species diversity and evenness were cula, Cymbella and Nitzschia in the Paisuni and Ken, observed to be 3.03 and 0.50, respectively, in the and Navicula, Nitzschia,andCymbella in the Tons, Betwa of the same ecoregion. The species diver- contribute to this trend (Appendix B). sity is higher in the Central Highlands ecoregion PCA and cluster analysis on diatom abundance ( HD 5.56) compared with the Himalayan (H D indicated longitudinal patterns in contrast to the 4.40) and other Plateau rivers (Nautiyal and Nau- spatial patterns (among the river sections). The flo- tiyal, 1999a). Biodiversity maximizes at interme- ral composition (species abundance) is similar in diate levels of disturbance (Townsend et al., 1987; the lowland sections of the Ken and Tons due to Minshall, 1988), which can operate on a local similar ecological conditions (soft sediments and scale. Factors like predation, herbivory, fluctua- flooding). However, other sections of the Ken are tions in physical factors, and catastrophes can be similar to the Paisuni, attributable to similar ripar- lumped together as disturbance (Krebs, 1994). ian conditions (Saal forest and prevalence of agri- Human activities also act as disturbances, indi- culture) in their basins. rectly in the form of agriculture and directly in the form of impoundments across the Betwa. Benthic macroinvertebrates The taxonomic composition of diatom flora from The macroinvertebrate fauna identified in our the Central Highlands is dominated by biraphids. In study is comparable with other Peninsular and

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Himalayan rivers/basins. No definite pattern of macroinvertebrate fauna declines generally from invertebrate richness is evident for the tropical or the headwaters to the lowland sections of each temperate conditions. The number of benthic river, except the Paisuni (Mishra, and Nautiyal, fauna is high at the regional scale rather than in 2011; Nautiyal and Mishra, 2012; Mishra and individual rivers because the species differ Nautiyal, 2013a). The macroinvertebrate fauna of between one site and the next (Allan, 1996; Vin- the Central Highlands consists largely of insects. son and Hawkins, 1998). The mollusc and annelid elements are very few in In the case of benthic macroinvertebrate fauna, the fauna. This is comparable with insect-domi- the taxonomic richness in the Ken and Paisuni is nated fauna from rivers of the Western Ghats relatively the same, though the Ken is longer than (Sivaramakrishnan et al., 1995; Subramanian and the Paisuni. The Tons, which is relatively shorter Sivaramkrishnan, 2005), as well as the mountain than the Ken, has the highest richness among all part of the Gangetic drainage in Uttarakhand. In three rivers. In these rivers, the richness does not contrast, mollusc-dominated fauna has been increase with river length (or area). Instead, the reported from the rivers of the Malwa Plateau higher richness in the Tons may be attributed to (Kulshrestha, 2009) in the western part of the Cen- the possibility of more habitat availability caused tral Highlands. by substrate heterogeneity in the Tons, a concept The most abundant macroinvertebrate taxon is accepted in island biogeography (Begon et al., longitudinally similar in various sections of the 1990). However, few studies are available indicat- rivers Ken, Paisuni, and Tons. Longitudinal varia- ing that richness increases with respect to sam- tions in abundance are attributed primarily to sub- pling site (Milesi et al., 2009; Nautiyal and strate preferences of the fauna and land-use Mishra, 2012). At a spatial scale, macroinverte- practices in the rivers of this ecoregion (Mishra brate richness increases slightly from the Ken to and Nautiyal, 2011). It can be argued that similar- the Tons in the lowland sections, while a reverse ity of substrate and riparian conditions along the trend of decrease is seen with regard to species river lead to longitudinal similarity of the fauna, as density. The lowland section of the Paisuni sup- it appears in the Paisuni and Tons but not in the ports high richness and density compared with the Ken, as was also observed by Nautiyal and Mishra lowland sections of the other two rivers. The rich- (2012). Vannote et al. (1980) state that gradients ness does not differ in the lowland sections of the of physical and chemical characteristics from Ken and Tons, where substrate heterogeneity is headwater to mouth result in different community greatly reduced by annual flooding that deposits and functional feeding groups (FFG). Longitudi- smaller and softer sediments on the river beds. nally, collectors are dominant in all the rivers. At latitudes comparable with the Central High- Therefore, the rivers are functionally heterotrophic land rivers, 58 taxa were recorded from the Brazil- (Mishra and Nautiyal, 2013b). This study is in ian Plateau rivers (27N) and 25 taxa were found agreement with the River Continuum Concept, as in the Lules River basin (26 N; Mesa, 2010). At heterotrophic conditions prevail in the headwater the tropical latitude (12N) of India, macroinverte- sections of the rivers (i.e. third order), though in brate richness (43 taxa) was relatively higher in the present study these rivers represent a third the River Kaveri (Sivaramakrishnan et al., 1995). order condition from headwater to mouth. Corkum (1992) observed that density was not con- The faunal abundance (shown as percent) also sistent among the rivers of different biomes. exhibits longitudinal similarity within each river The diversity of benthic macroinvertebrates is (intra-basin), as the dominant taxa are identical, known to increase with increasing substrate het- especially in the headwater and middle sections of erogeneity (Singh et al., 1994; Nautiyal and Mis- the Ken and Tons. However, in the lowland sec- hra, 2012) represented by the middle and lower tions of the Ken and Tons in the Gangetic Plains, sections of the Ken and Paisuni Rivers. Diversity the fauna lacks inter-basin similarity. This is is high in the agricultural lands (Richards et al. attributed to the deposition of soft sediments by 1997) and lower in urbanized areas (Walsh et al. the Yamuna and the Ganga around the confluence 2001). The proximate factors appear to be more of these rivers during annual flooding. Thus, the responsible for the assemblages of the benthic rivers are not similar section by section with macroinvertebrate fauna than large-scale factors respect to the macroinvertebrate fauna. Cluster (Strayer et al., 2003). The density of benthic analysis for the river sections clearly indicates that

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the faunal abundance in the headwater or middle are not similar section by section with respect to sections of the Ken and Tons are not similar, the macroinvertebrate fauna. The diatom taxa A. though the rivers originate in close proximity. chitrakootense is specifically abundant in the Pai- Only the lowland section of the Ken and Tons suni. Navicula seminulum and Navicula rostellata resemble one another. The headwater section of are characteristic of the Paisuni and the middle the Paisuni is similar to the headwater, middle, section of Tons. The Paisuni, which is three times and lower sections of the Tons, while the lower shorter than the Ken and Tons, is exceptional in and lowland sections of the Paisuni are similar to some respects, being unique both in respect to ben- the lower section of the Ken. Pan et al. (1999) thic diatom and macroinveretebrate species, while showed that the spatial patterns of diatom assemb- the Ken and Tons resemble each other relatively lages in the Mid-Atlantic Highlands were best dis- more. Though the longitudinal trends for species criminated by both land cover/land use in richness and density of benthic flora and fauna are catchments and by site-specific factors such as similar, the examined communities differ in ele- riparian conditions, as observed in the Central ments and abundance. Thus, generalisations Highlands ecoregion. regarding similar flora, fauna, and communities in the rivers of the ecoregion cannot be made for the Factors governing distributional patterns of Central Highland rivers. In other words, these riv- macroinvertebrates ers are diverse because the species differ between CCA indicated that land use, current velocity, one site and the next. In view of the Ken–Betwa and substratum are important variables for distri- river link, this ecoregion needs to be examined for bution, while depth and water temperature diversity of different trophic levels (NWDA, emerge as the most important factors with respect 2006). to the ecoregion. These factors have figured in Species richness of diatoms is high in the Cen- other studies too: current velocity in the Paisuni tral Highland rivers compared to temperate rivers (Mishra and Nautiyal, 2011), land use in the Ken (Kawecka and Olech, 2004). The floral composi- (Nautiyal and Mishra, 2012); substratum in the tion and abundance at subtropical latitudes in the Tons (Mishra and Nautiyal, 2013a), and current Himalayas (Nautiyal et al., 2013) is different from velocity, substrate size, conductivity, and abun- the Central Highland rivers. Species richness of dance of aquatic plants in the Patagonian Plateau macroinvertebrates in the Central Highland rivers rivers (Miserendino and Pizzolon, 2003). somewhat resembles that of the Plateau rivers of South America (Mesa, 2010), but is certainly The central highland rivers: Are they diverse? lower than that found in the tropical latitudes of Rivers of an ecoregion are expected to harbour India itself (Sivaramakrishnan et al., 1995). The similar flora, fauna, and communities. This faunal elements differ considerably from the Dec- hypothesis was examined in respect to the rich- can Plateau at tropical latitudes, as well as subtrop- ness, density, and taxonomic composition of ben- ical Plateau rivers and basins of South America thic diatom and macroinvertebrate assemblages in (Nautiyal and Mishra, 2012). three Plateau rivers. Observations reveal that the three rivers are similar in species richness of the Science and management diatom community, but the rivers differ in respect Study of the benthic community is important to species richness of benthic macroinvertebrate because it reflects changes in the aquatic environ- fauna. With some exceptions, the pattern of gen- ment either due to natural or anthropogenic eral decrease in the species richness and density effects. The Ken–Betwa river link has been from headwater to lowland sections is also similar approved to sustain and enhance agricultural pro- for both communities among these rivers. All riv- duction in the Bundelkhand region. The link ers support high diatom species diversity. The spe- involves building another dam on the Ken and cies-rich genera (Navicula, Cymbella and diverting the water to the Betwa through a linkage Nitzschia) were also similar. The insect fauna pre- canal (Figure 1). Presently, the fish fauna of the vails in stretches of river with hard substrate, while Betwa are threatened by habitat fragmentation mollusc and annelid species are found in lowland (Lakra et al., 2010). In addition to social impacts sections having soft sediments. The faunal ele- (Krueger et al., University of Michigan Master of ments lack inter-basin similarity. Thus, the rivers Science Degree, 2007), the link will reduce spatial

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heterogeneity, constrain the species represented at landings. Presently, Cyprinus carpio, Labeo cal- sites (Pan et al., 2000), and hinder its role as a vital basu, and miscellaneous groups constitute the junction for the mountain biotic elements of the major portion of capture fishery in the lower sec- subcontinent, especially in the donor river, Ken tions of the Ken, Paisuni and Tons. (NAAS, 2004, Policy Paper 28). The present study has demonstrated that the Plateau rivers are diverse with certain fish species being specific to Conclusions some rivers (Appendix E), as are some diatom species. Maintaining scientifically recommended In the present study, taxonomic richness and flows with regular monitoring is thus imperative density of benthic flora and fauna decrease longi- for the Plateau river ecosystems. tudinally from the headwater to lowland sections in all the rivers (except for the macroinvertebrate Priority gaps and needs species in the Paisuni). This is due to the gradual The present study explains only the Bundelk- reduction of substrate particle size from boulder hand part of the Central Highlands ecoregion, (stony) to silt-clay (soft), which provide low sur- while the rest remain unknown. It is important to face area for production of producer and consumer generate information on this ecoregion because of communities. The abundant benthic flora varies the native elements and the region’s value as a longitudinally in all the rivers, but in the case of vital junction of mountain biota. benthic fauna variation occurs only in the Ken, but there is similarity from middle to lowland in the Speculation on future trends and potential for Paisuni as well as to the headwater, middle and fish species, fisheries, and habitats lowland in the Tons. The Central Highlands ecore- The Betwa has more fish species (63) than the gion supports high richness and diversity of dia- Ken (56), the Paisuni (52), and the Tons (48). toms. Spatially, there is no definite trend in the Many (43) species are common to all three rivers. flora and fauna (except in the lowland sections) in The species richness of fish declines eastwards in all measured aspects (richness, density, and abun- the Central Highlands ecoregion. The food web dant taxa). Therefore, each river and section has comprising allochthonous C autocthonous inputs, its own characteristic flora and fauna, which sug- along with the producer community of which dia- gests that the proximate factors govern the various toms constitute a very significant part, and the features of assemblages, rather than regional fac- invertebrate consumer community that utilises tors such as latitude, altitude, and climate. This detritus and algal biomass, are the major drivers of study demonstrates high diversity across these fishery. Although, the species richness of benthic river basins in the Central Highlands ecoregion. communities declines in the rivers from source to mouth, the species richness of fish fauna increases in the lower part of the rivers as they widen, pro- Acknowledgements viding a variety of habitats that various fish species can utilise. Homogeneous softer sediments, Co-authors acknowledge the financial assistance decreased current velocity, and a greater volume for the Doctor of Philosophy program at the Uni- of water creates river depths that allow the plank- versity of Allahabad. We thank H.N.B Garhwal ton community to flourish. Because of this, there University, Srinagar for academic support. is a decline in benthic species richness. Sperata spp. is highly prized followed by L. rohita, T. tor, while L. calbasu is the least prized Supplemental material fish for commercial fisheries. The high exploita- tion rate for Tor tor in the Paisuni and Tons Riv- Supplemental data for this article can be ers, and L. rohita in the Ken has threatened these accessed on the publisher’s website. species (Dwivedi and Nautiyal, 2012). These rivers are stressed habitats because shallow river sections dry up during summer. The river linking will ORCID reduce desired flows, reduce fish diversity, and the more tolerant common carp will dominate the fish Prakash Nautiyal http://orcid.org/0000-0002-7958-5755

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