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Behavioural and Functioning Interactions in the Schizothoracid Community in the River Mandakini: An Assessment Through Altering Sex Ratio Patterns PDF

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Journal of the Bombay Natural History Society, 101 (3), Sep-Dec 2004 374-380 BEHAVIOURAL AND FUNCTIONING INTERACTIONS IN THE SCHIZOTHORACID COMMUNITY IN THE RIVER MANDARINE AN ASSESSMENT THROUGH ALTERING SEX RATIO PATTERNS' N. Singh2 3andK.C. Bhatt2 'Accepted November, 2001 'Zoology Department, P.0 Box 83, HNB Garhwal University, Srinagar Garhwal, Uttaranchal, India. 'Email: [email protected] Theinterrelationshipsandfunctioningofthreecloselyrelatedspecies,SchizothoraxplagiostomusHeckel,Schizothorax richardsonii Gray and Schizothoraichthys progastus McClelland (Teleostei, Family Cyprinidae, Subfamily Schizothoracinae), comprising the snow trout or schizothoracid community, as reflected by the altering sex ratio patterns, have been assessed quantitatively. Observations were made fortwo years, from January 1991 to December 1992,atfourlandingsites: Bheri(1,020m),Chandrapuri(827m),Agastyamuni(760m)andTilwara(724m),covering the lowerstretch ofover30 km ofthe high altitudeglacier-fed River Mandakini intheGarhwal Himalaya. The water characteristicsoftheriver(totalwaterdischarge,watervelocityandwatertemperature)weremeasuredatChandrapuri. Thetotalfishcatchofthesespeciesfromallthesiteswasfirstpooledandarrangedaccordingtosex,percentage,breeding and non-breeding seasons. The sex ratio was determined. These species breed twice ayear, from March to May and AugusttoOctober, with well-defined non-breedingintervals. Thesespeciesindividuallyfollowedasimilarpatternof responseto changes in the environment, as revealed by the altering sex ratio patterns i.e., highest values during first breedingseasonof1991 and 1992(2.65:1,3.0:1,3.89:1 and2.56:1,3.86:1,4.75:1 respectively)whenwaterparameters began rising from the lowest in the extreme winter. The lowest values ofsex ratio were observed during the second breedingseasonsof1991 and 1992(2.03:1, 1.51:1, 1.58:1 and1.59:1, 1.82:1, 1.74:1 respectively)whenwatercharacteristics valueswereatapeak,orjustbeganplummetingfromthepeaksofmonsoon.Thecorrespondingvaluesofsexratiointhe entireschizothoracid communitywereobservedas 2.78:1, 2.83:1 and 1.91:1, 1.61:1 respectively, duringthefirstand secondbreedingseasonsof1991, 1992. Sexratiobeginstoalteratthecommencementofjustrisingand/orpeak,orjust plummetingperiods.Alteringsexratiopattern isoneoftheindicatorsofthebeginningofthebreedingprocess.Thatis thefishtakethechangesin surroundingsasthecuetoinitiatebreedingprocess. Boththespeciesandthecommunityof schizothoracinefishesaltertheirsexratio pattern in responseto changes intheirsurroundings. Thispattern can serve asanindicatorofthefunctioningandinterrelationshipatindividual,population,speciesandcommunitylevels.Patterns ofsexratiochangemustbeutilizedforassessingthebehaviouralandevolutionaryprocessesoveraperiodaswellasan effective tool in the regulative management ofsnow trout community in the high altitude glacier-fed hillstreams of Garhwal Himalaya. Keywords: Schizothoracidcommunity,sexratio, RiverMandakini,functioninginteractions,behaviouralindicators, evolutionary processes INTRODUCTION herbivorous)andSchizothoraichthysprogastusMcClelland (columndwellerand feeder,carni-omnivorous)areanalysed Odum (1971) and Kendeigh (1980) described the asfurtherextensionofearlierworks(Bhatt 1993; Singh 1995, conceptual framework of biotic communities. In fluvial 1997; Singh and Subbaraj 2000; Singh etal. 1996)focusing systems, unidirectional flow and gradient determine the on the high altitude glacier-fed hill stream, Mandakini in mobility of the subject. This offers entirely different GarhwalHimalaya. challenges in the ways ofstudying the biotic communities Thesexratiooffishpopulation isaneffective indicator therein. The biotic communities in the rapidly flowing offunctioning, behavioural strategiesand catch composition hillstreams are heterogeneous assemblage of several plant (Nikolskii 1980;Engenwaji 1992). IntheIndiansubcontinent, andanimaltaxaofdiversefeatureswithintricaterelationships. with immensescopeofdiversityand potentialsoffish fauna, The upstream and downstream boundaries ofthe habitat(s) the assessmentoffunctioningand interrelationships through and movement of the residents for diverse purposes are sexratioand/orsimilarindicesoffishesatpopulation,species difficult to determine. In view of these restraints, the and community levelsremains unexplored, thus, eliminating behavioural attributes in terms ofaltering sex ratio patterns their advantageous uses for determining the characteristics, of schizothoracid or snowtrout community consisting of planning and management ofcultivable and non-cultivable Schizothorax plagiostomus Heckel, Schizothorax fishes to any worthwhile level (Pantulu 1961; Bhatt 1993; richardsonii Gray (both bottom dwellers and feeders, Nautiyal 1994). INTERACTIONS IN THE SCHIZOTHORACIDCOMMUNITY IN RIVER MANDAKINI The dominant status, economic potentials yet hitherto size, the percentage of males and females during the little explored avenues ofschizothoracid community, in the corresponding period suitably conformed to the respective entire Himalayan region, in general, and in Garhwal hills in structural representation in the community (Fig. 2). Further, particular, offer enough incentives for assessing the the sample size did not cast its shadow on the overall functioning interrelationships, their observable indices (sex response(s) patterns. ratiochangebeingoneofthese),andresponse(s)tothemilieu Percentage ofmales and females: The analysis offish of changing surroundings of glacier-fed hillstreams in the catchintennsofsexpercentageoftwoyearsofferedsimilarity region likeRiverMandakini. of patterns which the three species exhibited respectively (Table 1, Fig. 1). Forthe entire community, the corresponding MATERIALAND METHODS valuesofpercentageofmaleswere 9.31, 13.12and 12.83, 14.39 respectively during the first and second breeding season of The study is based on the analysis of the total fish 1991 and 1992. Forfemales the percentages were found to be catch of Schizothorax plagiostomus, Schizothorax 3.35,6.87and4.53,8.93 respectivelyduringthecorresponding richardsoniiand Schizothoraichthysprogastus in the lower breedingandnon-breedingseasonsof1991 and 1992(Fig.2). stretch of over 30 km of glacier-fed high altitude River Sex ratio: It is an effective and definite indicator of Mandakini, which finally meets with the still larger River response(s)and interactionswithinthecommunityaswell as Alaknanda at Rudraprayag (JBNHS 94(2), 1997: Fig. I, to the changing surroundings. The analysis samples of p.418).Therando—msamplingofadultspecimenswasmadeat Schizothorax plagiostomus, Schizothorax richardsonii and four landingsites Bheri (1,020 m), Chandrapuri (827 in), Schizothoraichthys progastus revealed a consistent pattern Agustmuni(760m),andTilwara(724m). Sinceallthesethree ofsexratioalterations(Table2,Fig. 1).Allthreespeciesshowed species breed twice a year, from March-May and August- higher sex ratio (2.78:1, 2.83:1 respectively) during the first October, the data were analysed and computed on the basis breeding season ofthe year but lower figures (1.91:1, 1.61:1 ofa pooled number as sample size, percentage ofmales and respectively)duringsecond breedingoftheyear(Fig. 2). femalesinthetotalcatch,breedingandnon-breedingseasons WaterparametersoftheRiverMandakini(Figs 1,2): with respect to individual species separately. The ratio of Thehabitatofthethreespeciesofschizothoracidcommunity M:F(M-numberofmales,F-numberoffemales)duringthe in the River Mandakini, when analysed and quantified for corresponding seasons gave the sex ratio ofeach species. total water discharge, water velocity and watertemperature, — The data ofwater parameters ofRiver Mandakini presented an overview conforming to the overall picture of total water discharge (m—Vsec), water velocity (m/sec) and what had been stated so far. The mean monthly values of water temperature (°C) during the corresponding study rivercharacteristicsbeganrisingfromMarch-May 1991, 1992 period have been obtained from the Uttar Pradesh Irrigation fromthe lowestfiguresduringextremewintersandremained — Department, Srinagar Garhwal as measured at Chandrapuri moderate totalwaterdischarge(36.96, 88.32mVsec),water (JBNHS94(2), 1997: Fig. 1, p.418),andwereconverted into velocity(0.—745, 0.686 m/sec) and watertemperature (1 1.05, mean monthly values ofrespective seasons. 1 1.61 °C) when males in the community comprised the highest representation. Conversely, during the second RESULTS breeding season in August- September, October 1991, 1992 these parameters were either at the peak or hadjust started Sample size: The study is based on analysis ofa total coiningdown from thepeakvaluesofmonsoon; i.e., 155.34, of5,587 adultspecimens. Examination ofthetotal fish catch 170.63 mVsec; 1.805, 1.919 m/sec and 15.05°, 15.8 °C from all thefourlandingsiteswhenpooledtogetherrevealed respectively when the sex ratio touched the corresponding evident variations. The highest representation of lowest values. Thus, the water parameters of the River Schizothoraxplagiostomus[total4,585(82.07%),males3,120 Mandakiniperfectlysimulateand stimulatethepictureofthe (81.95%),females 1,465 (82.30%)] placedthisspeciesonthe community response(s) as a whole when viewed along with topstatusnotonly inthe schizothoracid community, butalso the corresponding data. among all the fish species resident in River Mandakini. Schizothorax richardsonii also had a good representation DISCUSSION [total674(12.06%),males455(1 1.96%),females219(12.30%)| but Schizothoraichthys progastus happened to be poorly (a) Majorstructural features such as dominant species represented [total 328(5.87%), males 232(6.09%), females like forms or indicators, and (b) physical habitat of the 96(5.39%)](Fig. 1).Despitethisenormousvariationinsample community and functional attributes such as type of 1 Bombay Nat. Hist. Soc., 101 (3), Sep-Dec 2004 375 INTERACTIONS IN THE SCHIZOTHORACID COMMUNITY IN RIVER MANDAKINI (°C) temp Water velocity (m/sec) Water water /sec) discharge3 Total (m F) : (M ratio Sex sample years males of of two %) size the Total (inof sample years females of of two size %) the Total (inof sample numbers) years of size two (pooledof Total Fig 1 SexratioofSchizothoraxplagiostomus(SP), Schizothoraxrichardsonii(SR) and Schizothoraichthysprogastus(SPRO)inrelationtosamplesize,percentageoffemalesandmales andwaterparameters(totaldischarge, velocity&temp. duringbreedingandnonbreedingseasons ) of1991-1992intheriverMandakim 376 J. Bombay Nat. Hist. Soc., 101 (3), Sep-Dec 2004 INTERACTIONS IN THE SCHIZOTHORACIDCOMMUNITY IN RIVER MANDAKINI community,metabolismetc.areaconvenientbasisfornaming However,BhattandPathak(1992)haveexplainedafewspecies and classifying the community (Odum 1971). Of these, atcommunity level. functionalapproachesofferbetteralternativesforcomparison From quantitative and qualitative estimates ofpresent ofallcommunitiesinwidelydifferenthabitats. Regardingthe study, the topmost status of schizothoracid community is fish communities withinthe geographical regions, the major evidentamongvariousanimalcommunitiesinclusiveofthose approachesmustinclude-(1)Zonalapproachforrecognizing, ofotherfishcommunitiesresidentinRiverMandakini. Based classifying and listing in a sort ofcheck-list ofcommunity on dominance, various species ofschizothoracid community type, (2) gradient analysis approach involving the may be arranged, from higher to lower ranks, as arrangement ofpopulation along uni- or multi-dimensional (1) Schizothorax p/agiostomus, (2) Schizothorax environmental gradient axis with community recognition richardsonii (3) Schizothoraichthysprogastus (Singh et al. , based on statistical comparison. Also, there are certain 1996) and so on. The variation in the quantitative values of specifically interesting questions about fish communities - different functional attributes (including sex ratio, breeding (i) To what extent can fish in similar habitat be predicted? seasons)atdifferentsites isduetothecontinuumofcomplex (ii)Howarefishcommunitiesorganized?Doesthelastmember dynamics, altering interactions of individuals-individuals, ofthe sensitive species result in the collapse or irreversible individual-population, population-population, population- changes in the community? (iii) How is the state of species, species-species, species-community, community- development or completeness measured, and hence community and also with surroundings in each case and at assessment ofthe extent to which it has been degraded by each level apartfromworkingmodalities. environmental change(s). Notable examples ofsuch studies The highest absolute numbers of specimens during oftemperatefishcommunitiesofNorthAmericaincludethose breeding seasons, especially at Chandrapuri and Tilwara by Harvey, 1975, 1978; Johnson et al. 1977; Tonn and (,JBNHS 94(2), 1997: Fig. 1, p. 418, where smaller rivulets Magnuson 1983. debouch into river Mandakini), percentage of males and Theanalyticalstudiesoffishcommunitiesinthe Indian femalesinthetotalcatchduringMarch,April,May 1991, 1992 context are not addressed to in right perspectives despite and then during August, September, October, 1991, 1992 the sizable scope of diversity, functional changes within respectively, are convincing proofofthese species breeding shorter time and space (Shastri et al. 1982), potentials of twice a year (see Singh 1997). At the onset of breeding, newerdistinctapproacheslikelytoemerge. InGarhwalregion, potential male and female brooders migrate from lower Kumar(1991)analysedmicrozoobenthiccommunitiesofRiver stretches of larger glacier fed hill streams (like Ganga, Alaknanda, and Rawat (1992) undertook the analysis Alaknandaetc.)totheupperreachesoftheirtributaries. Small of community structure of plankton in high altitude Lake glacier/non-glacierfed streams orrivuletswith fasterwater DeoriaTal. velocity, lesserturbidityetc.aremoreconduciveenvironsfor The functional approach of community analysis of brooders,spawnandjuveniles. DestinationslikeChandrapuri Indian fishes in diverse fluvial systems wherein the subject andTilwara(sitesofconfluence)aremorepreferredsitesfor aswellasphysical habitathavehigh mobilityand, hence,the spawning and breeding purpose. community functioning and interactions are at continuous Singh (1997) mentioned several intrinsicand extrinsic — change areratherobscureobviouslydueto lowfeasibility. factors interacting together and collectively responsible for Table 1: Profile of pecentage of males and females ofthree species of schizothoracid community during breeding and non-breeding seasons of 1991-1992 in the River Mandakini Species Sex November-February March-May June-July August-October (1st Non-breeding) (1st breeding) (2nd Non-breeding) (2nd breeding) Jan-Feb Nov-Dec Jan-Feb Nov-Dec 1991 1992 1991 1992 1991 1992 1991 1991 1992 1992 Sp. 1 Male 5.03 9.68 3.01 12.72 19.07 3.09 5.26 19.10 22.18 Sp 2 Male 6.37 7.25 2.42 11.21 17.80 6.15 7.69 23.52 17.58 Sp 3 Male 7.76 5.60 3.45 31 89 16.38 6.03 1.72 12.93 14.22 Sp 1 Female 4.70 7.92 2.73 10.24 15.77 3.75 5.19 20.00 2969 Sp 2 Female 6.85 7.76 2.74 7.76 9.59 5.94 6.85 32.42 20 09 Sp 3 Female 10.42 9.38 4.16 19.79 8.33 6.25 2.08 19.79 19.79 Sp. 1 = Schizothorax plagiostomus, Sp. 2 = Schizothorax richardsonii. Sp 3 = Schizothoraichthys progastus J. Bombay Nat. Hist. Soc., 101 (3), Sep-Dec 2004 377 INTERACTIONS IN THE SCHIZOTHORACIDCOMMUNITY IN RIVER MANDAKINI (°C) temp Water velocity (m/sec) Water water /sec) discharge3 Total (m F) : (M ratio Sex years sampleand of males two of size %) Total (infemales numbers, seperately) (pooled years females two sample of of and size males Total Fig. 2: SexratioofSchizothoracidcommunityinrelationtopoolednumbersaswellasmales andfemalesseparately, %offemales&malesandwaterparameters(totaldischarge, velocity&temp. duringbreedingand ) non-breedingseasonsof1991-1992intheRiverMandakini(ComparewithFig. 1) 378 J. Bombay Nat. Hist. Soc., 101 (3), Sep-Dec 2004 INTERACTIONS IN THE SCHIZOTHORACID COMMUNITY IN RIVER MANDAKINI Table 2 Profile ofthe sex ratios ofthree species of schizothoracid community during breeding and non-breeding seasons of 1991-1992 in the River Mandakini Species November-February March-May June-July August-October (1st Non-breeding) (1st breeding) (2nd Non-breeding) (2nd breeding) Jan-Feb Nov-Dec Jan-Feb Nov-Dec 1991 1992 1991 1992 1991 1992 1991 1991 1992 1992 Sp 1 2.28:1 2.60:1 2.35:1 2.65:1 2.56:1 2.24:1 2.16:1 2.03:1 1.59:1 Sp 2 1.93:1 1.94:1 1.83:1 3.00:1 3.86:1 2.16:1 2.33:1 1.51:1 1.82:1 Sp 3 1.80:1 3.67:1 2.00:1 3.89:1 4.75:1 2.33:1 2.00:1 1.58:1 1.74:1 Schizothoracid 2.17:1 2.38:1 2.26:1 2.82:1 2.79:1 2.23:1 2.18:1 1.91:1 1.61:1 Community Sp 1 = Schizothorax plagiostomus, Sp. 2 = Schizothorax richardsonii, Sp 3 = Schizothoraichthys progastus higher male sex ratio during the first breeding season ofthe inhabitingsimilarhabitatandusingsimilarcuesforspawning, year: (i)earlierdepartureofmalesand latearrival offemales, breedingand other vital processes? Here lie the possibilities (ii)freshrecruitmentofnewbatchesofsubadultsintobrooders, of preponement and/or postponement of such changes in (iii) difficulties encountered by female brooders during behavioural indicesand indicators(likesexratioalterations) upstream migration because of their full-grown belly and by a few hours/days enough for such resource mobilization, relatively lowerwaterdischargetocopewiththe largersized resource partitioning and minimizing the competition for females.Theconditionsotherwiseareconduciveforspawning resources at successive levels within the community which and breeding resulting from moderate conditions, and musthave been too intense. It isalso indicative ofintelligent (iv) vulnerabilities offemale brooderstotheir predators and designing of behavioural strategies of schizothoracids in othernaturalhazards.Also,environmentalconditionsduring natureformobilizationoftransitional butmoderateresources the first breeding season favour the male, whereas females in terms oftimingand space for breeding. are favoured during the second breeding period ofthe year. This analysis of altering sex ratio patterns in Another possibility of sex ratio alterations may be sex Schizothorax plagiostomus Schizothorax richardsonii and , dependentmortalityandintersexes in population(anentirely Schizothoraichthysprogastus presents an overall picture in newdimension). the River Mandakini in particular and other glacier-fed Since the present assessment is based on the data of hillstreams of Garhwal region in general. This study of threecloselyrelatedspeciesofschizothoracids, itindicatesa schizothoracid community, though raising several questions similar picture of response(s) as evident from sex ratio tobeansweredfromvariouspointsofview,shouldbeutilized variations in relation to changing river characteristics. This in two ways; first, in revealingthe processes ofsignallingof similaritylendsfurthercredencetotheirclosegenetickinship individual-population-species-community responses, astheyinhabitandsharecommonhabitat(theriverMandakini convergence-divergence at behavioural, physiological and andlikesintheHimalayanregion)andalsoencountersimilar environmental levels, and second, in regulative management challengesandopportunities, usesimilarcuesandstimulants. ofschizothoracidfisheryinglacier-fedhighaltitudehillstreams Obviously, they must have evolved similar behavioural ofGarhwalregion. repertoire, functioning strategies and response(s) at individual,population,speciesandcommunitylevels.Varying ACKNOWLEDGEMENTS sex ratio patterns, mobilization ofspawning grounds (Singh 1995) are among these. Such behavioural responses ofthe The author is grateful to Prof. AshaChandola Saklani, community also include the interplay of intra- and inter- Head and Co-ordinator, UGC-SAP Department ofZoology, specific convergence, divergence/diversity, which are at the HNBGarhwalUniversity, SrinagarGarhwalforherconsistent helms ofevolutionary processes. Sex ratio alterations also support and for being available for critical suggestions at reflect the pattern diversity of stratification (Pielou 1966), successive stages throughout the study tenure. Financial zonation, activity, reproduction, social, co-action, assistance received from the Government ofIndia, Ministry stochasticism etc. However, many significant questions ofEnvironment,ForestsandWildlife,NewDelhi(No. 35/20/ emanate out ofsuch a scenario, namely levels ofintra- and 90-RE) (presently monitored by GBPIHED, Kosi, Almora, inter-specific convergence and divergence/diversity while Uttaranchal) is thankfully acknowledged. J. Bombay Nat. Hist. Soc., 101 (3), Sep-Dec 2004 379 INTERACTIONS INTHE SCHIZOTHORACID COMMUNITY IN RIVER MANDAKINI REFERENCES Bhatt, K.C. (1993): Present status and olfaction ofsnowtrout in the Co., London. Reprinted byNatraj Publishers, Dehra Dun, 1996. river Mandakini (Garhwal Himalaya). Ph.D. dissertation, HNB Pp. 145-161. Garhwal University, SrinagarGarhwal,India. Pp. 1-152. Pantulu, V.R. (1961): Determination ofage and growth ofMystus Bhatt, S.D. & J.K. Pathak (1992): Himalayan Environment: Water gulio Ham. byuseofpectoral spine in observation initsbiology Quality ofDrainage Basins. Shri Almora Book Depot, Almora. and fisheries in the Houghly estuary. Proc. Nat. Inst. India Sci. Pp. 1-318. B22: 4. Engenwaji, H.M.G. (1992): Reproductive Biology of four African Pielou, E.C. (1966): Species diversity and pattern diversity in the catfishes(Osteichthys:Claridae)inAnambrariverbasin,Nigeria. studyofecological succession.J. Theoret. Biol. 10: 370-383. Hydrobiologia 242: 155-164. 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Bahuguna (1996): Schizothoracids in Nikolskii,GV.(1980):Theoryoffishpopulationdynamics,(translated the river Mandakini ofGarhwal Himalaya: a briefnote oftheir by J.E.S. Brodley, Ed.: Jones, R.), Bishen Singh Mahendra Pal status. J. Anim. Morphol Physiol. 43(2): 163-168. Singh,DehraDun&OttoKoeltzSciencePublishing,Koenigesten, Tonn, W.M. & J.J. Magnuson (1983): Communityanalysis in Fishery Germany. Pp. 88-138. management: an application with Wisconsin Lakes. Trans. Fish. Odum, E.P. (1971): FundamentalsofEcology. 3rded . W.B. Saunders Soc. 112: 377-3 86. 380 J. Bombay Nat. Hist. Soc., 101 (3), Sep-Dec 2004

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