7
Research Article Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch, 1793) from Northern and Eastern Regions of India Using Truss Analysis Ankur Kashyap, Madhu Awasthi, and Mohammad Serajuddin Department of Zoology, University of Lucknow, Lucknow, Uttar Pradesh 226007, India Correspondence should be addressed to Mohammad Serajuddin; lu.fi[email protected] Received 31 October 2015; Accepted 13 January 2016 Academic Editor: Michel Laurin Copyright © 2016 Ankur Kashyap et al. is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Morphometric variation study was carried out using truss analysis in freshwater murrel, Channa punctatus (Bloch, 1793), to explore different phenotypic stocks of C. punctatus. Significant variation in the morphometric analysis was observed using the truss network and multivariate analysis of the three subpopulations of freshwater murrel, C. punctatus, from the two different regions of India. Principal component analysis accounted for a total variation of 91.95% by the first three principal components. PC 1 represented the variation due to size and PC 2 and PC 3 represented variations due to shape. e 95% confidence ellipses depicted the clear isolation between the samples in scatter graph of PC 1 on PC 2. e combined groups plot (DF 1 on DF 2) based on discriminant function analysis (DFA) also validated the existence of three different subpopulations. In group classification using DFA, 100 percent of all the samples were correctly classified into their original subpopulation. ese results support the existence of multiracial composition of C. punctatus across the different geographical isolated sites and, thus, these findings indicate the presence of three different phenotypic stocks of C. punctatus. 1. Introduction Stock identification is considered to be an important aspect in the field of fishery management because the stocks with variable life history attributes such as growth and reproduc- tive potentials are considered to be important to increase yield and also for stock improvement programme of fish. Variation in the growth, development, and maturation in different members of the same species results in variety of body shapes [1]. Patterns of morphometric variation in fishes indicate differences in growth and maturation rates because the body forms are products of ontogeny. e multiple stock compositions of the fish were reported by many workers [2– 4]. e truss network system was used by many notable workers to segregate the stocks [5–7]. e truss analysis is important because it covers the entire shape of the fish in a form of a uniform network, and thus the probability of extracting the morphometric differences among the different fish populations is increased. Working on Indian fishes like Catla catla, Rastrelliger kanagurta, and Megalaspis cordyla,a limited number of workers used the truss morphometry to discriminate the stocks [8–10]. A survey of literature showed that no such work on morphometric variations using truss was carried out on C. punctatus so far, except for the work carried out by Khan et al. [11]. erefore, the present study was carried out to find out the morphometric variations employing truss network system using modern geometric morphometric methods to segregate the phenotypic stocks of C. punctatus collected from different isolated geographical sites. 2. Materials and Methods e total of 90 fish samples were collected from three different regions, that is, Gomti River at Lucknow ( = 30, 26 52 N 80 55 E) and ponds situated at Kolkata ( = 30, 22 34 N 88 22 E) and Malihabad ( = 30, 26 55 N 80 43 E). ese regions of the country are geographically isolated Hindawi Publishing Corporation International Journal of Zoology Volume 2016, Article ID 2605404, 6 pages http://dx.doi.org/10.1155/2016/2605404

Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

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Page 1: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

Research ArticlePhenotypic Variation in Freshwater MurrelChanna punctatus (Bloch 1793) from Northern andEastern Regions of India Using Truss Analysis

Ankur Kashyap Madhu Awasthi and Mohammad Serajuddin

Department of Zoology University of Lucknow Lucknow Uttar Pradesh 226007 India

Correspondence should be addressed to Mohammad Serajuddin lufisheriesgmailcom

Received 31 October 2015 Accepted 13 January 2016

Academic Editor Michel Laurin

Copyright copy 2016 Ankur Kashyap et alThis is an open access article distributed under the Creative Commons Attribution Licensewhich permits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

Morphometric variation study was carried out using truss analysis in freshwater murrelChanna punctatus (Bloch 1793) to exploredifferent phenotypic stocks ofC punctatus Significant variation in themorphometric analysis was observed using the truss networkand multivariate analysis of the three subpopulations of freshwater murrel C punctatus from the two different regions of IndiaPrincipal component analysis accounted for a total variation of 9195 by the first three principal components PC 1 represented thevariation due to size and PC 2 and PC 3 represented variations due to shapeThe 95 confidence ellipses depicted the clear isolationbetween the samples in scatter graph of PC 1 on PC 2 The combined groups plot (DF 1 on DF 2) based on discriminant functionanalysis (DFA) also validated the existence of three different subpopulations In group classification usingDFA 100 percent of all thesamples were correctly classified into their original subpopulation These results support the existence of multiracial compositionof C punctatus across the different geographical isolated sites and thus these findings indicate the presence of three differentphenotypic stocks of C punctatus

1 Introduction

Stock identification is considered to be an important aspectin the field of fishery management because the stocks withvariable life history attributes such as growth and reproduc-tive potentials are considered to be important to increaseyield and also for stock improvement programme of fishVariation in the growth development and maturation indifferent members of the same species results in variety ofbody shapes [1] Patterns of morphometric variation in fishesindicate differences in growth and maturation rates becausethe body forms are products of ontogeny The multiple stockcompositions of the fish were reported by many workers [2ndash4]

The truss network system was used by many notableworkers to segregate the stocks [5ndash7] The truss analysis isimportant because it covers the entire shape of the fish ina form of a uniform network and thus the probability ofextracting the morphometric differences among the differentfish populations is increased Working on Indian fishes like

Catla catla Rastrelliger kanagurta and Megalaspis cordyla alimited number of workers used the truss morphometry todiscriminate the stocks [8ndash10] A survey of literature showedthat no such work on morphometric variations using trusswas carried out on C punctatus so far except for the workcarried out by Khan et al [11] Therefore the present studywas carried out to find out the morphometric variationsemploying truss network system using modern geometricmorphometric methods to segregate the phenotypic stocksof C punctatus collected from different isolated geographicalsites

2 Materials and Methods

Thetotal of 90 fish sampleswere collected from three differentregions that is Gomti River at Lucknow (119899 = 30 26∘ 521015840N 80∘ 551015840 E) and ponds situated at Kolkata (119899 = 30 22∘341015840 N 88∘ 221015840 E) and Malihabad (119899 = 30 26∘ 551015840 N 80∘ 431015840E) These regions of the country are geographically isolated

Hindawi Publishing CorporationInternational Journal of ZoologyVolume 2016 Article ID 2605404 6 pageshttpdxdoiorg10115520162605404

2 International Journal of Zoology

Figure 1 Location of 11 morphometric landmarks for constructingthe truss network system on Channa punctatus

from each other Images of specimens of C punctatus weretaken using digital camera (Nikon 160 megapixels) A totalof eleven morphometric landmarks (Figure 1) were digitizedin software TpsDig version 140 [12] For covering the entireshape of fish two-dimensional procrustes fitting was donewith software Paleontological Statistics [13] after TpsDig and23 truss measurements were taken for morphometric studyas depicted in Figure 1 Prior to data analysis the size effectwas removed from the data as the collected samples of fishwere not of the same age and size Significant correlationswere observed between the size (standard length) and themorphometric truss measurements So 119872 transformationand then log transformation of all measurements was carriedout to remove the size effect and normalization of the datarespectivelyThe size effect resulting was eliminated from thedata using following formula [14]

119872adj = 119872 lowast (119871 119904119871119900)119887

(1)

where119872 is original measurement119872adj is size-adjusted mea-surement 119871119900 is standard length 119871 119904 is the mean of standardlength and parameter 119887 was estimated for each characterfrom observed data as the slope of regression of log119872 onlog 119871119900 using all fish in all populations The 119872 transformeddata was again checked for correlations of morphometricmeasurements with standard length for efficiency check ofsize removal Standard length and total length were excludedfrom the final analysis Both univariate and multivariateanalysis of variance were carried out to test the significanceof morphological variations

After data transformation all the truss measurementswere subjected to one-way ANOVA analysis to find out thesignificant (119901 lt 05) morphometric truss measurementsbetween the subpopulations of C punctatus Significant (119901 lt05) morphometric measurements were further subjectedto principal component analysis (PCA) and discriminantfunction analysis (DFA) to interpret the size and shapevariation and for group classification respectively All thecalculations were done with help of PAST and SPSS 120software

3 Results and Discussion

None of the 23 truss measurements gave significant cor-relation with the standard length after 119872 transformation

Table 1 Highly significant (119901 lt 001) 23 truss measurementsbetween the three subpopulations of C punctatus in one-wayANOVA analysis

S number Morphometric measurements 119901 value1 1-2 0002 1ndash10 0003 1ndash11 0004 2-3 0005 2ndash9 0006 2ndash10 0007 2ndash11 0008 3-4 0009 3ndash8 00010 3ndash9 00011 3ndash10 00012 4-5 00013 4ndash7 00014 4ndash8 00015 4ndash9 00016 5ndash7 00017 5ndash8 00018 5-6 00019 7-8 00020 6-7 00021 8-9 00022 9-10 00023 10-11 000

indicating that allometric formula was effective in removingsize effect from the data All the 23 truss measurementswere found to be highly significant (119901 lt 001) in one-wayANOVA (Table 1) and were further tested in multivariateanalysis using PCA and DFA

The principal component analysis resulted in extractionof 23 principal components The first three componentsextracted accounted for a total variance of 8325 533 and337 by PC 1 PC 2 and PC 3 respectively Three separate95 confidence ellipses were obtained in PC 1 scatter graphon PC 2 (Figure 2) The most significant loadings on PC 1were nearly same in magnitude and found to be positivelycorrelated to all the measurements (Table 2) The PC 2 andPC 3 were positively and negatively correlated to all themeasurements The high components loadings of PC 2 weretruss measurements 4-5 10-11 2ndash10 1ndash10 and 1ndash11 and thosecorrelated with PC 3 were 4-5 10-11 9-10 3ndash10 2ndash10 and 1ndash10(Figures 3 and 4) These measurements were mainly respon-sible for the variation between the three subpopulationsIn discriminant function analysis (DFA) 100 of originalgrouped cases were correctly classified into their respectivesubpopulation and two discriminant functions were formed(DF 1 and DF 2) The two functions accounted for 100variability between the groups in which DF 1 and DF 2explained 996 and 04 of the between-group variationfor morphometric measurements The combined group plotof DF 1 on DF 2 indicated the existence of three different

International Journal of Zoology 3

Table 2 First three principal components extracted from principalcomponent analysis

S number Morphometricmeasurements PC 1 PC 2 PC 3

1 1-2 08985 minus01621 015892 1ndash10 09459 02164 minus019283 1ndash11 09282 minus0185 0012244 2-3 0904 01695 00096575 2ndash9 09589 01141 0088926 2ndash10 09227 03027 minus019947 2ndash11 09732 minus007009 012618 3-4 09515 01206 017129 3ndash8 09661 006335 0112710 3ndash9 09344 00342 0124211 3ndash10 09196 minus009022 021512 4-5 07589 minus05035 minus0388713 4ndash7 09687 minus01191 minus00966514 4ndash8 09677 minus004644 00370715 4ndash9 0954 007266 0125516 5ndash7 09918 00384 00142417 5ndash8 09837 minus002992 00346518 5-6 09507 01036 00274819 7-8 08632 minus01474 minus00285520 6-7 08704 01604 00888921 8-9 09461 001748 00512522 9-10 08742 minus01059 0318823 10-11 07943 04871 minus03151

Total variance () 8325 533 337Cumulative variance

() 8325 8858 9195

phenotypic stocks of C punctatus as three isolated groupcentroids were obtained in DFA (Figure 5) Examinationof the pooled within-group correlations of discriminatingfunctions and morphometric measurements revealed thatmeasurements 5ndash7 3ndash8 5ndash8 2ndash9 4ndash9 4ndash8 8-9 3ndash9 3ndash10 1ndash10 and 9-10 were mainly responsible for differences amongthe subpopulations (Table 3)

The present truss study revealed the high degree ofsignificant morphometric heterogeneity among the subpop-ulations of C punctatus The univariate (one-way ANOVA)and multivariate analysis via PCA and DFA suggested threephenotypic distinct stocks ofC punctatusThesemorphome-tric differences can be attributed to the fact that these threesubpopulations of C punctatus are geographically isolatedfrom each other

Principal component analysis laid emphasis on the effectof size on the linear dimensions of themeasurements apart forexplaining the shape variation in the three subpopulationsPC 1 (8325 variance) was positively correlated to all themorphometric traits represented by the general size but PC2 and PC 3 explained about 533 and 337 variancerespectively due to shape as they were strongly positively and

0 16 32 48 64Component 1

0

08

16

24

Com

pone

nt 2

minus08

minus16

minus24

minus32

minus4

minus16minus32minus48minus64minus8

Figure 2 Scatter plot of PC 1 on PC 2 showing 95 confidenceellipses of three subpopulations of C punctatus Red violet andblue colours indicate Gomti River pond of Malihabad and pond ofKolkata subpopulation respectively

Table 3 Pooled within-groups correlations between discriminatingmorphometric measurements and standardized canonical discrim-inant functions Variables ordered by absolute size of correlationwithin function

S number Morphometric measurements Function 1 Function 21 5ndash7 173lowast 0222 3ndash8 152lowast minus0853 5ndash8 136lowast 0614 2ndash9 117lowast minus0945 4ndash9 114lowast minus0986 4ndash8 094lowast minus0137 8-9 092lowast minus0228 3ndash9 083lowast 0079 3ndash10 072lowast 06110 1ndash10 071lowast 02111 9-10 058lowast 05212 1-2 072 362lowast

13 6-7 064 minus301lowast14 2ndash11 135 277lowast

15 4-5 033 235lowast

16 3-4 134 minus160lowast17 1ndash11 067 159lowast

18 2-3 065 minus148lowast19 5-6 093 minus144lowast20 4ndash7 086 113lowast

21 7-8 047 113lowast

22 2ndash10 064 minus087lowast23 10-11 036 minus066lowastlowastLargest absolute correlation between each variable and any discriminantfunction

4 International Journal of Zoology

0216401695

01141

03027

01206

00633500342

00726600384

01036

01604

001748

04871

A B C D E F G H I J K L M N O P Q R S T U V W

0

012

024

036

048C

orre

latio

n

minus012

minus024

minus036

minus048

minus06

minus01059

minus01474

minus002992minus004644

minus01191

minus009022minus007009

minus0185

minus01621

minus05035

Figure 3 PCA loadings of PC 2 and its correlation with various morphometric parameters

01589

0012240009657

008892

01261

01712

0112701242

0215

003707

01255

001424003465

002748

008889 005125

03188

A B C D E F G H I J K L M N O P Q R S T U V W

0

008

016

024

032

Cor

rela

tion

minus008

minus016

minus024

minus032

minus04

minus03151

minus002855

minus009665

minus01994minus01928

minus03887

Figure 4 PCA loadings of PC 3 and its correlation with various morphometric parameters

negatively correlated to morphometric traitsThe coefficientsof same sign in PCs and coefficients with negative andpositive signs of any section of PCs are indicative of ldquosizevariationrdquo and ldquoshape variationrdquo respectively [15ndash18]

The consistent differences in the morphometric mea-surements between the three subpopulations of C punctatussuggested the restricted intermingling among the subpop-ulations of Lucknow Malihabad and Kolkata which wasalso supported by the nonoverlapping and isolated groupcentroids in DFA Saila and Flowers [19] used DFA to studythe geographic variation by analyzing the morphometrics ofAmerican lobster The existence of different strains of Indian

major carp Catla catla was reported by NBFGR [20] andemphasized that it was due to complete geographical isolationwithout any intermixing The morphometric variations inthe different subpopulations of C punctatus in the presentstudy may be due to geographical isolation and differentenvironmental conditions However it is generally assumedthat these differences may be genetically related or might beassociated with phenotypic plasticity in response to differentenvironmental factors of various habitats [21] Moreoverthe variations recorded among three subpopulations of Cpunctatus could be a result of phenotypic plasticity expressedin response to the environmental conditions The present

International Journal of Zoology 5

minus50 minus25 0 25 50

Function 1

Func

tion2

Canonical discriminant functions

PopulationsGomti River at LucknowPond of Kolkata

Pond of MalihabadGroup centroid

50

25

0

minus25

minus50

1

23

Figure 5 Group centroids of the three subpopulations of Cpunctatus in DFA combined groups plot

results are similar to those of Pathak et al [22] who reportedmorphometric variation inMacrognathus pancalus from twodifferent river basins of India Khan et al [11] also discrim-inated the subpopulations of C punctatus collected fromthree different rivers (Ganga Yamuna and Gomti) of Indiawhere the differences were in the head abdomen and caudalpeduncle regions The major contributions to DFA and PCAwere the measurements belonging to these regions includingthe high allometric shape contributions from diagonal bodymeasurements

4 Conclusion

These current findings in the truss analysis indicated signifi-cant morphometric variations and thus the presence of threedifferent phenotypic stocks of C punctatus

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors thank the Head of the Department of ZoologyUniversity of Lucknow for providing facility and one ofthe authors is thankful to the Department of Science andTechnology (DST-PURSE) for providing him fellowship

References

[1] S X Cadrin ldquoAdvances in morphometric identification offishery stocksrdquo Reviews in Fish Biology and Fisheries vol 10 no1 pp 91ndash112 2000

[2] A Kashyap M Awasthi and M Serajuddin ldquoIntraspecificdiversity of freshwatermurrelChanna punctatusrdquo InternationalJournal of Fisheries andAquatic Studies vol 3 no 2 pp 166ndash1732015

[3] J Verma A Kashyap and M Serajuddin ldquoPhylogeny basedon truss analysis in five populations of freshwater catfishClupisoma garuardquo International Journal of Science and Researchvol 3 no 8 pp 1414ndash1418 2014

[4] A Kashyap M Awasthi and M Serajuddin ldquoGeographicmorphometric variations of freshwater murrel Channa punc-tatus from northern and eastern parts of Indiardquo Proceedingsof the National Academy of Sciences India Section B BiologicalSciences pp 1ndash7 2014

[5] L Hauser G R Carvalho and T J Pitcher ldquoMorphologicaland genetic differentiation of the African clupeid Limnothrissamiodon 34 years after its introduction to Lake Kivurdquo Journal ofFish Biology vol 47 pp 127ndash144 1995

[6] D G Bembo G R Carvalho N Cingolani E Arneri GGiannetti and T J Pitcher ldquoAllozymic and morphometricevidence for two stocks of the European anchovy Engraulisencrasicolus in Adriatic watersrdquo Marine Biology vol 126 no 3pp 529ndash538 1996

[7] C Turan ldquoA note on the examination of morphometric dif-ferentiation among fish populations the truss systemrdquo TurkishJournal of Zoology vol 23 pp 259ndash264 1999

[8] N C Ujjainia and M P S Kohli ldquoLandmark-based morpho-metric analysis for selected species of Indian major carp (Catlacatla Ham 1822)rdquo International Journal of Food Agriculture andVeterinary Science vol 1 pp 64ndash74 2011

[9] P Jayasankar P CThomasM P Paulton and J Mathew ldquoMor-phometric and genetic analyzes of Indian mackerel (Rastrelligerkanagurta) from peninsular Indiardquo Asian Fisheries Science vol17 pp 201ndash215 2004

[10] A M Sajina S K Chakraborty A K Jaiswar D GPazhayamadam and D Sudheesan ldquoStock structure analysisof Megalaspis cordyla (Linnaeus 1758) along the Indian coastbased on truss network analysisrdquo Fisheries Research vol 108 no1 pp 100ndash105 2011

[11] M A Khan K Miyan and S Khan ldquoMorphometric variationof snakehead fish Channa punctatus populations from threeIndian riversrdquo Journal of Applied Ichthyology vol 29 no 3 pp637ndash642 2013

[12] F J Rholf TpsDig Version 140 Department of Ecology andEvolution State University of New York New York NY USA2004

[13] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysisrdquo Palaeontologia Electronica vol 4 article 9 2001

[14] N G Elliott K Haskard and J A Koslow ldquoMorphometricanalysis of orange roughy (Hoplostethus atlanticus) off thecontinental slope of southern Australiardquo Journal of Fish Biologyvol 46 no 2 pp 202ndash220 1995

[15] P Jolicoeur and J EMosimann ldquoSize and shape variation in thepainted turtle A principal component analysisrdquoGrowth vol 24pp 339ndash354 1960

[16] J M Humphries F L Bookstein B Chernoff G R Smith R LElder and S G Poss ldquoMultivariate discrimination by shape in

6 International Journal of Zoology

relation to sizerdquo Systematic Biology vol 30 no 3 pp 291ndash3081981

[17] F L Bookstein ldquolsquoSize and Shapersquo a comment on semanticsrdquoSystematic Biology vol 38 no 2 pp 173ndash180 1989

[18] P Sundberg ldquoShape and size-constrained principal componentsanalysisrdquo Systematic Zoology vol 38 no 2 pp 166ndash168 1989

[19] S B Saila and J M Flowers ldquoGeographic morphometricvariation in the American lobsterrdquo Systematic Zoology vol 18no 3 pp 330ndash338 1969

[20] National Bureau of Fish Genetic Resources Annual Report ofNational Bureau of Fish Genetic Research National Bureau ofFish Genetic Resources Lucknow India 2001

[21] A G Murta ldquoMorphological variation of horse mackerel (Tra-churus trachurus) in the Iberian and North African Atlanticimplications for stock identificationrdquo ICES Journal of MarineScience vol 57 no 4 pp 1240ndash1248 2000

[22] B C Pathak J I Mir andM Serajuddin ldquoMorphometric varia-tion among barred spiny eelMacrognathus pancalus (Hamilton1822) populations from the Ganges and Brahmaputra riverbasin India by using geomorphometricsrdquo Journal of Biologyvol 3 no 1 pp 15ndash20 2013

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

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BioinformaticsAdvances in

Marine BiologyJournal of

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Signal TransductionJournal of

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International Journal of

Microbiology

Page 2: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

2 International Journal of Zoology

Figure 1 Location of 11 morphometric landmarks for constructingthe truss network system on Channa punctatus

from each other Images of specimens of C punctatus weretaken using digital camera (Nikon 160 megapixels) A totalof eleven morphometric landmarks (Figure 1) were digitizedin software TpsDig version 140 [12] For covering the entireshape of fish two-dimensional procrustes fitting was donewith software Paleontological Statistics [13] after TpsDig and23 truss measurements were taken for morphometric studyas depicted in Figure 1 Prior to data analysis the size effectwas removed from the data as the collected samples of fishwere not of the same age and size Significant correlationswere observed between the size (standard length) and themorphometric truss measurements So 119872 transformationand then log transformation of all measurements was carriedout to remove the size effect and normalization of the datarespectivelyThe size effect resulting was eliminated from thedata using following formula [14]

119872adj = 119872 lowast (119871 119904119871119900)119887

(1)

where119872 is original measurement119872adj is size-adjusted mea-surement 119871119900 is standard length 119871 119904 is the mean of standardlength and parameter 119887 was estimated for each characterfrom observed data as the slope of regression of log119872 onlog 119871119900 using all fish in all populations The 119872 transformeddata was again checked for correlations of morphometricmeasurements with standard length for efficiency check ofsize removal Standard length and total length were excludedfrom the final analysis Both univariate and multivariateanalysis of variance were carried out to test the significanceof morphological variations

After data transformation all the truss measurementswere subjected to one-way ANOVA analysis to find out thesignificant (119901 lt 05) morphometric truss measurementsbetween the subpopulations of C punctatus Significant (119901 lt05) morphometric measurements were further subjectedto principal component analysis (PCA) and discriminantfunction analysis (DFA) to interpret the size and shapevariation and for group classification respectively All thecalculations were done with help of PAST and SPSS 120software

3 Results and Discussion

None of the 23 truss measurements gave significant cor-relation with the standard length after 119872 transformation

Table 1 Highly significant (119901 lt 001) 23 truss measurementsbetween the three subpopulations of C punctatus in one-wayANOVA analysis

S number Morphometric measurements 119901 value1 1-2 0002 1ndash10 0003 1ndash11 0004 2-3 0005 2ndash9 0006 2ndash10 0007 2ndash11 0008 3-4 0009 3ndash8 00010 3ndash9 00011 3ndash10 00012 4-5 00013 4ndash7 00014 4ndash8 00015 4ndash9 00016 5ndash7 00017 5ndash8 00018 5-6 00019 7-8 00020 6-7 00021 8-9 00022 9-10 00023 10-11 000

indicating that allometric formula was effective in removingsize effect from the data All the 23 truss measurementswere found to be highly significant (119901 lt 001) in one-wayANOVA (Table 1) and were further tested in multivariateanalysis using PCA and DFA

The principal component analysis resulted in extractionof 23 principal components The first three componentsextracted accounted for a total variance of 8325 533 and337 by PC 1 PC 2 and PC 3 respectively Three separate95 confidence ellipses were obtained in PC 1 scatter graphon PC 2 (Figure 2) The most significant loadings on PC 1were nearly same in magnitude and found to be positivelycorrelated to all the measurements (Table 2) The PC 2 andPC 3 were positively and negatively correlated to all themeasurements The high components loadings of PC 2 weretruss measurements 4-5 10-11 2ndash10 1ndash10 and 1ndash11 and thosecorrelated with PC 3 were 4-5 10-11 9-10 3ndash10 2ndash10 and 1ndash10(Figures 3 and 4) These measurements were mainly respon-sible for the variation between the three subpopulationsIn discriminant function analysis (DFA) 100 of originalgrouped cases were correctly classified into their respectivesubpopulation and two discriminant functions were formed(DF 1 and DF 2) The two functions accounted for 100variability between the groups in which DF 1 and DF 2explained 996 and 04 of the between-group variationfor morphometric measurements The combined group plotof DF 1 on DF 2 indicated the existence of three different

International Journal of Zoology 3

Table 2 First three principal components extracted from principalcomponent analysis

S number Morphometricmeasurements PC 1 PC 2 PC 3

1 1-2 08985 minus01621 015892 1ndash10 09459 02164 minus019283 1ndash11 09282 minus0185 0012244 2-3 0904 01695 00096575 2ndash9 09589 01141 0088926 2ndash10 09227 03027 minus019947 2ndash11 09732 minus007009 012618 3-4 09515 01206 017129 3ndash8 09661 006335 0112710 3ndash9 09344 00342 0124211 3ndash10 09196 minus009022 021512 4-5 07589 minus05035 minus0388713 4ndash7 09687 minus01191 minus00966514 4ndash8 09677 minus004644 00370715 4ndash9 0954 007266 0125516 5ndash7 09918 00384 00142417 5ndash8 09837 minus002992 00346518 5-6 09507 01036 00274819 7-8 08632 minus01474 minus00285520 6-7 08704 01604 00888921 8-9 09461 001748 00512522 9-10 08742 minus01059 0318823 10-11 07943 04871 minus03151

Total variance () 8325 533 337Cumulative variance

() 8325 8858 9195

phenotypic stocks of C punctatus as three isolated groupcentroids were obtained in DFA (Figure 5) Examinationof the pooled within-group correlations of discriminatingfunctions and morphometric measurements revealed thatmeasurements 5ndash7 3ndash8 5ndash8 2ndash9 4ndash9 4ndash8 8-9 3ndash9 3ndash10 1ndash10 and 9-10 were mainly responsible for differences amongthe subpopulations (Table 3)

The present truss study revealed the high degree ofsignificant morphometric heterogeneity among the subpop-ulations of C punctatus The univariate (one-way ANOVA)and multivariate analysis via PCA and DFA suggested threephenotypic distinct stocks ofC punctatusThesemorphome-tric differences can be attributed to the fact that these threesubpopulations of C punctatus are geographically isolatedfrom each other

Principal component analysis laid emphasis on the effectof size on the linear dimensions of themeasurements apart forexplaining the shape variation in the three subpopulationsPC 1 (8325 variance) was positively correlated to all themorphometric traits represented by the general size but PC2 and PC 3 explained about 533 and 337 variancerespectively due to shape as they were strongly positively and

0 16 32 48 64Component 1

0

08

16

24

Com

pone

nt 2

minus08

minus16

minus24

minus32

minus4

minus16minus32minus48minus64minus8

Figure 2 Scatter plot of PC 1 on PC 2 showing 95 confidenceellipses of three subpopulations of C punctatus Red violet andblue colours indicate Gomti River pond of Malihabad and pond ofKolkata subpopulation respectively

Table 3 Pooled within-groups correlations between discriminatingmorphometric measurements and standardized canonical discrim-inant functions Variables ordered by absolute size of correlationwithin function

S number Morphometric measurements Function 1 Function 21 5ndash7 173lowast 0222 3ndash8 152lowast minus0853 5ndash8 136lowast 0614 2ndash9 117lowast minus0945 4ndash9 114lowast minus0986 4ndash8 094lowast minus0137 8-9 092lowast minus0228 3ndash9 083lowast 0079 3ndash10 072lowast 06110 1ndash10 071lowast 02111 9-10 058lowast 05212 1-2 072 362lowast

13 6-7 064 minus301lowast14 2ndash11 135 277lowast

15 4-5 033 235lowast

16 3-4 134 minus160lowast17 1ndash11 067 159lowast

18 2-3 065 minus148lowast19 5-6 093 minus144lowast20 4ndash7 086 113lowast

21 7-8 047 113lowast

22 2ndash10 064 minus087lowast23 10-11 036 minus066lowastlowastLargest absolute correlation between each variable and any discriminantfunction

4 International Journal of Zoology

0216401695

01141

03027

01206

00633500342

00726600384

01036

01604

001748

04871

A B C D E F G H I J K L M N O P Q R S T U V W

0

012

024

036

048C

orre

latio

n

minus012

minus024

minus036

minus048

minus06

minus01059

minus01474

minus002992minus004644

minus01191

minus009022minus007009

minus0185

minus01621

minus05035

Figure 3 PCA loadings of PC 2 and its correlation with various morphometric parameters

01589

0012240009657

008892

01261

01712

0112701242

0215

003707

01255

001424003465

002748

008889 005125

03188

A B C D E F G H I J K L M N O P Q R S T U V W

0

008

016

024

032

Cor

rela

tion

minus008

minus016

minus024

minus032

minus04

minus03151

minus002855

minus009665

minus01994minus01928

minus03887

Figure 4 PCA loadings of PC 3 and its correlation with various morphometric parameters

negatively correlated to morphometric traitsThe coefficientsof same sign in PCs and coefficients with negative andpositive signs of any section of PCs are indicative of ldquosizevariationrdquo and ldquoshape variationrdquo respectively [15ndash18]

The consistent differences in the morphometric mea-surements between the three subpopulations of C punctatussuggested the restricted intermingling among the subpop-ulations of Lucknow Malihabad and Kolkata which wasalso supported by the nonoverlapping and isolated groupcentroids in DFA Saila and Flowers [19] used DFA to studythe geographic variation by analyzing the morphometrics ofAmerican lobster The existence of different strains of Indian

major carp Catla catla was reported by NBFGR [20] andemphasized that it was due to complete geographical isolationwithout any intermixing The morphometric variations inthe different subpopulations of C punctatus in the presentstudy may be due to geographical isolation and differentenvironmental conditions However it is generally assumedthat these differences may be genetically related or might beassociated with phenotypic plasticity in response to differentenvironmental factors of various habitats [21] Moreoverthe variations recorded among three subpopulations of Cpunctatus could be a result of phenotypic plasticity expressedin response to the environmental conditions The present

International Journal of Zoology 5

minus50 minus25 0 25 50

Function 1

Func

tion2

Canonical discriminant functions

PopulationsGomti River at LucknowPond of Kolkata

Pond of MalihabadGroup centroid

50

25

0

minus25

minus50

1

23

Figure 5 Group centroids of the three subpopulations of Cpunctatus in DFA combined groups plot

results are similar to those of Pathak et al [22] who reportedmorphometric variation inMacrognathus pancalus from twodifferent river basins of India Khan et al [11] also discrim-inated the subpopulations of C punctatus collected fromthree different rivers (Ganga Yamuna and Gomti) of Indiawhere the differences were in the head abdomen and caudalpeduncle regions The major contributions to DFA and PCAwere the measurements belonging to these regions includingthe high allometric shape contributions from diagonal bodymeasurements

4 Conclusion

These current findings in the truss analysis indicated signifi-cant morphometric variations and thus the presence of threedifferent phenotypic stocks of C punctatus

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors thank the Head of the Department of ZoologyUniversity of Lucknow for providing facility and one ofthe authors is thankful to the Department of Science andTechnology (DST-PURSE) for providing him fellowship

References

[1] S X Cadrin ldquoAdvances in morphometric identification offishery stocksrdquo Reviews in Fish Biology and Fisheries vol 10 no1 pp 91ndash112 2000

[2] A Kashyap M Awasthi and M Serajuddin ldquoIntraspecificdiversity of freshwatermurrelChanna punctatusrdquo InternationalJournal of Fisheries andAquatic Studies vol 3 no 2 pp 166ndash1732015

[3] J Verma A Kashyap and M Serajuddin ldquoPhylogeny basedon truss analysis in five populations of freshwater catfishClupisoma garuardquo International Journal of Science and Researchvol 3 no 8 pp 1414ndash1418 2014

[4] A Kashyap M Awasthi and M Serajuddin ldquoGeographicmorphometric variations of freshwater murrel Channa punc-tatus from northern and eastern parts of Indiardquo Proceedingsof the National Academy of Sciences India Section B BiologicalSciences pp 1ndash7 2014

[5] L Hauser G R Carvalho and T J Pitcher ldquoMorphologicaland genetic differentiation of the African clupeid Limnothrissamiodon 34 years after its introduction to Lake Kivurdquo Journal ofFish Biology vol 47 pp 127ndash144 1995

[6] D G Bembo G R Carvalho N Cingolani E Arneri GGiannetti and T J Pitcher ldquoAllozymic and morphometricevidence for two stocks of the European anchovy Engraulisencrasicolus in Adriatic watersrdquo Marine Biology vol 126 no 3pp 529ndash538 1996

[7] C Turan ldquoA note on the examination of morphometric dif-ferentiation among fish populations the truss systemrdquo TurkishJournal of Zoology vol 23 pp 259ndash264 1999

[8] N C Ujjainia and M P S Kohli ldquoLandmark-based morpho-metric analysis for selected species of Indian major carp (Catlacatla Ham 1822)rdquo International Journal of Food Agriculture andVeterinary Science vol 1 pp 64ndash74 2011

[9] P Jayasankar P CThomasM P Paulton and J Mathew ldquoMor-phometric and genetic analyzes of Indian mackerel (Rastrelligerkanagurta) from peninsular Indiardquo Asian Fisheries Science vol17 pp 201ndash215 2004

[10] A M Sajina S K Chakraborty A K Jaiswar D GPazhayamadam and D Sudheesan ldquoStock structure analysisof Megalaspis cordyla (Linnaeus 1758) along the Indian coastbased on truss network analysisrdquo Fisheries Research vol 108 no1 pp 100ndash105 2011

[11] M A Khan K Miyan and S Khan ldquoMorphometric variationof snakehead fish Channa punctatus populations from threeIndian riversrdquo Journal of Applied Ichthyology vol 29 no 3 pp637ndash642 2013

[12] F J Rholf TpsDig Version 140 Department of Ecology andEvolution State University of New York New York NY USA2004

[13] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysisrdquo Palaeontologia Electronica vol 4 article 9 2001

[14] N G Elliott K Haskard and J A Koslow ldquoMorphometricanalysis of orange roughy (Hoplostethus atlanticus) off thecontinental slope of southern Australiardquo Journal of Fish Biologyvol 46 no 2 pp 202ndash220 1995

[15] P Jolicoeur and J EMosimann ldquoSize and shape variation in thepainted turtle A principal component analysisrdquoGrowth vol 24pp 339ndash354 1960

[16] J M Humphries F L Bookstein B Chernoff G R Smith R LElder and S G Poss ldquoMultivariate discrimination by shape in

6 International Journal of Zoology

relation to sizerdquo Systematic Biology vol 30 no 3 pp 291ndash3081981

[17] F L Bookstein ldquolsquoSize and Shapersquo a comment on semanticsrdquoSystematic Biology vol 38 no 2 pp 173ndash180 1989

[18] P Sundberg ldquoShape and size-constrained principal componentsanalysisrdquo Systematic Zoology vol 38 no 2 pp 166ndash168 1989

[19] S B Saila and J M Flowers ldquoGeographic morphometricvariation in the American lobsterrdquo Systematic Zoology vol 18no 3 pp 330ndash338 1969

[20] National Bureau of Fish Genetic Resources Annual Report ofNational Bureau of Fish Genetic Research National Bureau ofFish Genetic Resources Lucknow India 2001

[21] A G Murta ldquoMorphological variation of horse mackerel (Tra-churus trachurus) in the Iberian and North African Atlanticimplications for stock identificationrdquo ICES Journal of MarineScience vol 57 no 4 pp 1240ndash1248 2000

[22] B C Pathak J I Mir andM Serajuddin ldquoMorphometric varia-tion among barred spiny eelMacrognathus pancalus (Hamilton1822) populations from the Ganges and Brahmaputra riverbasin India by using geomorphometricsrdquo Journal of Biologyvol 3 no 1 pp 15ndash20 2013

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 3: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

International Journal of Zoology 3

Table 2 First three principal components extracted from principalcomponent analysis

S number Morphometricmeasurements PC 1 PC 2 PC 3

1 1-2 08985 minus01621 015892 1ndash10 09459 02164 minus019283 1ndash11 09282 minus0185 0012244 2-3 0904 01695 00096575 2ndash9 09589 01141 0088926 2ndash10 09227 03027 minus019947 2ndash11 09732 minus007009 012618 3-4 09515 01206 017129 3ndash8 09661 006335 0112710 3ndash9 09344 00342 0124211 3ndash10 09196 minus009022 021512 4-5 07589 minus05035 minus0388713 4ndash7 09687 minus01191 minus00966514 4ndash8 09677 minus004644 00370715 4ndash9 0954 007266 0125516 5ndash7 09918 00384 00142417 5ndash8 09837 minus002992 00346518 5-6 09507 01036 00274819 7-8 08632 minus01474 minus00285520 6-7 08704 01604 00888921 8-9 09461 001748 00512522 9-10 08742 minus01059 0318823 10-11 07943 04871 minus03151

Total variance () 8325 533 337Cumulative variance

() 8325 8858 9195

phenotypic stocks of C punctatus as three isolated groupcentroids were obtained in DFA (Figure 5) Examinationof the pooled within-group correlations of discriminatingfunctions and morphometric measurements revealed thatmeasurements 5ndash7 3ndash8 5ndash8 2ndash9 4ndash9 4ndash8 8-9 3ndash9 3ndash10 1ndash10 and 9-10 were mainly responsible for differences amongthe subpopulations (Table 3)

The present truss study revealed the high degree ofsignificant morphometric heterogeneity among the subpop-ulations of C punctatus The univariate (one-way ANOVA)and multivariate analysis via PCA and DFA suggested threephenotypic distinct stocks ofC punctatusThesemorphome-tric differences can be attributed to the fact that these threesubpopulations of C punctatus are geographically isolatedfrom each other

Principal component analysis laid emphasis on the effectof size on the linear dimensions of themeasurements apart forexplaining the shape variation in the three subpopulationsPC 1 (8325 variance) was positively correlated to all themorphometric traits represented by the general size but PC2 and PC 3 explained about 533 and 337 variancerespectively due to shape as they were strongly positively and

0 16 32 48 64Component 1

0

08

16

24

Com

pone

nt 2

minus08

minus16

minus24

minus32

minus4

minus16minus32minus48minus64minus8

Figure 2 Scatter plot of PC 1 on PC 2 showing 95 confidenceellipses of three subpopulations of C punctatus Red violet andblue colours indicate Gomti River pond of Malihabad and pond ofKolkata subpopulation respectively

Table 3 Pooled within-groups correlations between discriminatingmorphometric measurements and standardized canonical discrim-inant functions Variables ordered by absolute size of correlationwithin function

S number Morphometric measurements Function 1 Function 21 5ndash7 173lowast 0222 3ndash8 152lowast minus0853 5ndash8 136lowast 0614 2ndash9 117lowast minus0945 4ndash9 114lowast minus0986 4ndash8 094lowast minus0137 8-9 092lowast minus0228 3ndash9 083lowast 0079 3ndash10 072lowast 06110 1ndash10 071lowast 02111 9-10 058lowast 05212 1-2 072 362lowast

13 6-7 064 minus301lowast14 2ndash11 135 277lowast

15 4-5 033 235lowast

16 3-4 134 minus160lowast17 1ndash11 067 159lowast

18 2-3 065 minus148lowast19 5-6 093 minus144lowast20 4ndash7 086 113lowast

21 7-8 047 113lowast

22 2ndash10 064 minus087lowast23 10-11 036 minus066lowastlowastLargest absolute correlation between each variable and any discriminantfunction

4 International Journal of Zoology

0216401695

01141

03027

01206

00633500342

00726600384

01036

01604

001748

04871

A B C D E F G H I J K L M N O P Q R S T U V W

0

012

024

036

048C

orre

latio

n

minus012

minus024

minus036

minus048

minus06

minus01059

minus01474

minus002992minus004644

minus01191

minus009022minus007009

minus0185

minus01621

minus05035

Figure 3 PCA loadings of PC 2 and its correlation with various morphometric parameters

01589

0012240009657

008892

01261

01712

0112701242

0215

003707

01255

001424003465

002748

008889 005125

03188

A B C D E F G H I J K L M N O P Q R S T U V W

0

008

016

024

032

Cor

rela

tion

minus008

minus016

minus024

minus032

minus04

minus03151

minus002855

minus009665

minus01994minus01928

minus03887

Figure 4 PCA loadings of PC 3 and its correlation with various morphometric parameters

negatively correlated to morphometric traitsThe coefficientsof same sign in PCs and coefficients with negative andpositive signs of any section of PCs are indicative of ldquosizevariationrdquo and ldquoshape variationrdquo respectively [15ndash18]

The consistent differences in the morphometric mea-surements between the three subpopulations of C punctatussuggested the restricted intermingling among the subpop-ulations of Lucknow Malihabad and Kolkata which wasalso supported by the nonoverlapping and isolated groupcentroids in DFA Saila and Flowers [19] used DFA to studythe geographic variation by analyzing the morphometrics ofAmerican lobster The existence of different strains of Indian

major carp Catla catla was reported by NBFGR [20] andemphasized that it was due to complete geographical isolationwithout any intermixing The morphometric variations inthe different subpopulations of C punctatus in the presentstudy may be due to geographical isolation and differentenvironmental conditions However it is generally assumedthat these differences may be genetically related or might beassociated with phenotypic plasticity in response to differentenvironmental factors of various habitats [21] Moreoverthe variations recorded among three subpopulations of Cpunctatus could be a result of phenotypic plasticity expressedin response to the environmental conditions The present

International Journal of Zoology 5

minus50 minus25 0 25 50

Function 1

Func

tion2

Canonical discriminant functions

PopulationsGomti River at LucknowPond of Kolkata

Pond of MalihabadGroup centroid

50

25

0

minus25

minus50

1

23

Figure 5 Group centroids of the three subpopulations of Cpunctatus in DFA combined groups plot

results are similar to those of Pathak et al [22] who reportedmorphometric variation inMacrognathus pancalus from twodifferent river basins of India Khan et al [11] also discrim-inated the subpopulations of C punctatus collected fromthree different rivers (Ganga Yamuna and Gomti) of Indiawhere the differences were in the head abdomen and caudalpeduncle regions The major contributions to DFA and PCAwere the measurements belonging to these regions includingthe high allometric shape contributions from diagonal bodymeasurements

4 Conclusion

These current findings in the truss analysis indicated signifi-cant morphometric variations and thus the presence of threedifferent phenotypic stocks of C punctatus

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors thank the Head of the Department of ZoologyUniversity of Lucknow for providing facility and one ofthe authors is thankful to the Department of Science andTechnology (DST-PURSE) for providing him fellowship

References

[1] S X Cadrin ldquoAdvances in morphometric identification offishery stocksrdquo Reviews in Fish Biology and Fisheries vol 10 no1 pp 91ndash112 2000

[2] A Kashyap M Awasthi and M Serajuddin ldquoIntraspecificdiversity of freshwatermurrelChanna punctatusrdquo InternationalJournal of Fisheries andAquatic Studies vol 3 no 2 pp 166ndash1732015

[3] J Verma A Kashyap and M Serajuddin ldquoPhylogeny basedon truss analysis in five populations of freshwater catfishClupisoma garuardquo International Journal of Science and Researchvol 3 no 8 pp 1414ndash1418 2014

[4] A Kashyap M Awasthi and M Serajuddin ldquoGeographicmorphometric variations of freshwater murrel Channa punc-tatus from northern and eastern parts of Indiardquo Proceedingsof the National Academy of Sciences India Section B BiologicalSciences pp 1ndash7 2014

[5] L Hauser G R Carvalho and T J Pitcher ldquoMorphologicaland genetic differentiation of the African clupeid Limnothrissamiodon 34 years after its introduction to Lake Kivurdquo Journal ofFish Biology vol 47 pp 127ndash144 1995

[6] D G Bembo G R Carvalho N Cingolani E Arneri GGiannetti and T J Pitcher ldquoAllozymic and morphometricevidence for two stocks of the European anchovy Engraulisencrasicolus in Adriatic watersrdquo Marine Biology vol 126 no 3pp 529ndash538 1996

[7] C Turan ldquoA note on the examination of morphometric dif-ferentiation among fish populations the truss systemrdquo TurkishJournal of Zoology vol 23 pp 259ndash264 1999

[8] N C Ujjainia and M P S Kohli ldquoLandmark-based morpho-metric analysis for selected species of Indian major carp (Catlacatla Ham 1822)rdquo International Journal of Food Agriculture andVeterinary Science vol 1 pp 64ndash74 2011

[9] P Jayasankar P CThomasM P Paulton and J Mathew ldquoMor-phometric and genetic analyzes of Indian mackerel (Rastrelligerkanagurta) from peninsular Indiardquo Asian Fisheries Science vol17 pp 201ndash215 2004

[10] A M Sajina S K Chakraborty A K Jaiswar D GPazhayamadam and D Sudheesan ldquoStock structure analysisof Megalaspis cordyla (Linnaeus 1758) along the Indian coastbased on truss network analysisrdquo Fisheries Research vol 108 no1 pp 100ndash105 2011

[11] M A Khan K Miyan and S Khan ldquoMorphometric variationof snakehead fish Channa punctatus populations from threeIndian riversrdquo Journal of Applied Ichthyology vol 29 no 3 pp637ndash642 2013

[12] F J Rholf TpsDig Version 140 Department of Ecology andEvolution State University of New York New York NY USA2004

[13] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysisrdquo Palaeontologia Electronica vol 4 article 9 2001

[14] N G Elliott K Haskard and J A Koslow ldquoMorphometricanalysis of orange roughy (Hoplostethus atlanticus) off thecontinental slope of southern Australiardquo Journal of Fish Biologyvol 46 no 2 pp 202ndash220 1995

[15] P Jolicoeur and J EMosimann ldquoSize and shape variation in thepainted turtle A principal component analysisrdquoGrowth vol 24pp 339ndash354 1960

[16] J M Humphries F L Bookstein B Chernoff G R Smith R LElder and S G Poss ldquoMultivariate discrimination by shape in

6 International Journal of Zoology

relation to sizerdquo Systematic Biology vol 30 no 3 pp 291ndash3081981

[17] F L Bookstein ldquolsquoSize and Shapersquo a comment on semanticsrdquoSystematic Biology vol 38 no 2 pp 173ndash180 1989

[18] P Sundberg ldquoShape and size-constrained principal componentsanalysisrdquo Systematic Zoology vol 38 no 2 pp 166ndash168 1989

[19] S B Saila and J M Flowers ldquoGeographic morphometricvariation in the American lobsterrdquo Systematic Zoology vol 18no 3 pp 330ndash338 1969

[20] National Bureau of Fish Genetic Resources Annual Report ofNational Bureau of Fish Genetic Research National Bureau ofFish Genetic Resources Lucknow India 2001

[21] A G Murta ldquoMorphological variation of horse mackerel (Tra-churus trachurus) in the Iberian and North African Atlanticimplications for stock identificationrdquo ICES Journal of MarineScience vol 57 no 4 pp 1240ndash1248 2000

[22] B C Pathak J I Mir andM Serajuddin ldquoMorphometric varia-tion among barred spiny eelMacrognathus pancalus (Hamilton1822) populations from the Ganges and Brahmaputra riverbasin India by using geomorphometricsrdquo Journal of Biologyvol 3 no 1 pp 15ndash20 2013

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 4: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

4 International Journal of Zoology

0216401695

01141

03027

01206

00633500342

00726600384

01036

01604

001748

04871

A B C D E F G H I J K L M N O P Q R S T U V W

0

012

024

036

048C

orre

latio

n

minus012

minus024

minus036

minus048

minus06

minus01059

minus01474

minus002992minus004644

minus01191

minus009022minus007009

minus0185

minus01621

minus05035

Figure 3 PCA loadings of PC 2 and its correlation with various morphometric parameters

01589

0012240009657

008892

01261

01712

0112701242

0215

003707

01255

001424003465

002748

008889 005125

03188

A B C D E F G H I J K L M N O P Q R S T U V W

0

008

016

024

032

Cor

rela

tion

minus008

minus016

minus024

minus032

minus04

minus03151

minus002855

minus009665

minus01994minus01928

minus03887

Figure 4 PCA loadings of PC 3 and its correlation with various morphometric parameters

negatively correlated to morphometric traitsThe coefficientsof same sign in PCs and coefficients with negative andpositive signs of any section of PCs are indicative of ldquosizevariationrdquo and ldquoshape variationrdquo respectively [15ndash18]

The consistent differences in the morphometric mea-surements between the three subpopulations of C punctatussuggested the restricted intermingling among the subpop-ulations of Lucknow Malihabad and Kolkata which wasalso supported by the nonoverlapping and isolated groupcentroids in DFA Saila and Flowers [19] used DFA to studythe geographic variation by analyzing the morphometrics ofAmerican lobster The existence of different strains of Indian

major carp Catla catla was reported by NBFGR [20] andemphasized that it was due to complete geographical isolationwithout any intermixing The morphometric variations inthe different subpopulations of C punctatus in the presentstudy may be due to geographical isolation and differentenvironmental conditions However it is generally assumedthat these differences may be genetically related or might beassociated with phenotypic plasticity in response to differentenvironmental factors of various habitats [21] Moreoverthe variations recorded among three subpopulations of Cpunctatus could be a result of phenotypic plasticity expressedin response to the environmental conditions The present

International Journal of Zoology 5

minus50 minus25 0 25 50

Function 1

Func

tion2

Canonical discriminant functions

PopulationsGomti River at LucknowPond of Kolkata

Pond of MalihabadGroup centroid

50

25

0

minus25

minus50

1

23

Figure 5 Group centroids of the three subpopulations of Cpunctatus in DFA combined groups plot

results are similar to those of Pathak et al [22] who reportedmorphometric variation inMacrognathus pancalus from twodifferent river basins of India Khan et al [11] also discrim-inated the subpopulations of C punctatus collected fromthree different rivers (Ganga Yamuna and Gomti) of Indiawhere the differences were in the head abdomen and caudalpeduncle regions The major contributions to DFA and PCAwere the measurements belonging to these regions includingthe high allometric shape contributions from diagonal bodymeasurements

4 Conclusion

These current findings in the truss analysis indicated signifi-cant morphometric variations and thus the presence of threedifferent phenotypic stocks of C punctatus

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors thank the Head of the Department of ZoologyUniversity of Lucknow for providing facility and one ofthe authors is thankful to the Department of Science andTechnology (DST-PURSE) for providing him fellowship

References

[1] S X Cadrin ldquoAdvances in morphometric identification offishery stocksrdquo Reviews in Fish Biology and Fisheries vol 10 no1 pp 91ndash112 2000

[2] A Kashyap M Awasthi and M Serajuddin ldquoIntraspecificdiversity of freshwatermurrelChanna punctatusrdquo InternationalJournal of Fisheries andAquatic Studies vol 3 no 2 pp 166ndash1732015

[3] J Verma A Kashyap and M Serajuddin ldquoPhylogeny basedon truss analysis in five populations of freshwater catfishClupisoma garuardquo International Journal of Science and Researchvol 3 no 8 pp 1414ndash1418 2014

[4] A Kashyap M Awasthi and M Serajuddin ldquoGeographicmorphometric variations of freshwater murrel Channa punc-tatus from northern and eastern parts of Indiardquo Proceedingsof the National Academy of Sciences India Section B BiologicalSciences pp 1ndash7 2014

[5] L Hauser G R Carvalho and T J Pitcher ldquoMorphologicaland genetic differentiation of the African clupeid Limnothrissamiodon 34 years after its introduction to Lake Kivurdquo Journal ofFish Biology vol 47 pp 127ndash144 1995

[6] D G Bembo G R Carvalho N Cingolani E Arneri GGiannetti and T J Pitcher ldquoAllozymic and morphometricevidence for two stocks of the European anchovy Engraulisencrasicolus in Adriatic watersrdquo Marine Biology vol 126 no 3pp 529ndash538 1996

[7] C Turan ldquoA note on the examination of morphometric dif-ferentiation among fish populations the truss systemrdquo TurkishJournal of Zoology vol 23 pp 259ndash264 1999

[8] N C Ujjainia and M P S Kohli ldquoLandmark-based morpho-metric analysis for selected species of Indian major carp (Catlacatla Ham 1822)rdquo International Journal of Food Agriculture andVeterinary Science vol 1 pp 64ndash74 2011

[9] P Jayasankar P CThomasM P Paulton and J Mathew ldquoMor-phometric and genetic analyzes of Indian mackerel (Rastrelligerkanagurta) from peninsular Indiardquo Asian Fisheries Science vol17 pp 201ndash215 2004

[10] A M Sajina S K Chakraborty A K Jaiswar D GPazhayamadam and D Sudheesan ldquoStock structure analysisof Megalaspis cordyla (Linnaeus 1758) along the Indian coastbased on truss network analysisrdquo Fisheries Research vol 108 no1 pp 100ndash105 2011

[11] M A Khan K Miyan and S Khan ldquoMorphometric variationof snakehead fish Channa punctatus populations from threeIndian riversrdquo Journal of Applied Ichthyology vol 29 no 3 pp637ndash642 2013

[12] F J Rholf TpsDig Version 140 Department of Ecology andEvolution State University of New York New York NY USA2004

[13] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysisrdquo Palaeontologia Electronica vol 4 article 9 2001

[14] N G Elliott K Haskard and J A Koslow ldquoMorphometricanalysis of orange roughy (Hoplostethus atlanticus) off thecontinental slope of southern Australiardquo Journal of Fish Biologyvol 46 no 2 pp 202ndash220 1995

[15] P Jolicoeur and J EMosimann ldquoSize and shape variation in thepainted turtle A principal component analysisrdquoGrowth vol 24pp 339ndash354 1960

[16] J M Humphries F L Bookstein B Chernoff G R Smith R LElder and S G Poss ldquoMultivariate discrimination by shape in

6 International Journal of Zoology

relation to sizerdquo Systematic Biology vol 30 no 3 pp 291ndash3081981

[17] F L Bookstein ldquolsquoSize and Shapersquo a comment on semanticsrdquoSystematic Biology vol 38 no 2 pp 173ndash180 1989

[18] P Sundberg ldquoShape and size-constrained principal componentsanalysisrdquo Systematic Zoology vol 38 no 2 pp 166ndash168 1989

[19] S B Saila and J M Flowers ldquoGeographic morphometricvariation in the American lobsterrdquo Systematic Zoology vol 18no 3 pp 330ndash338 1969

[20] National Bureau of Fish Genetic Resources Annual Report ofNational Bureau of Fish Genetic Research National Bureau ofFish Genetic Resources Lucknow India 2001

[21] A G Murta ldquoMorphological variation of horse mackerel (Tra-churus trachurus) in the Iberian and North African Atlanticimplications for stock identificationrdquo ICES Journal of MarineScience vol 57 no 4 pp 1240ndash1248 2000

[22] B C Pathak J I Mir andM Serajuddin ldquoMorphometric varia-tion among barred spiny eelMacrognathus pancalus (Hamilton1822) populations from the Ganges and Brahmaputra riverbasin India by using geomorphometricsrdquo Journal of Biologyvol 3 no 1 pp 15ndash20 2013

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 5: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

International Journal of Zoology 5

minus50 minus25 0 25 50

Function 1

Func

tion2

Canonical discriminant functions

PopulationsGomti River at LucknowPond of Kolkata

Pond of MalihabadGroup centroid

50

25

0

minus25

minus50

1

23

Figure 5 Group centroids of the three subpopulations of Cpunctatus in DFA combined groups plot

results are similar to those of Pathak et al [22] who reportedmorphometric variation inMacrognathus pancalus from twodifferent river basins of India Khan et al [11] also discrim-inated the subpopulations of C punctatus collected fromthree different rivers (Ganga Yamuna and Gomti) of Indiawhere the differences were in the head abdomen and caudalpeduncle regions The major contributions to DFA and PCAwere the measurements belonging to these regions includingthe high allometric shape contributions from diagonal bodymeasurements

4 Conclusion

These current findings in the truss analysis indicated signifi-cant morphometric variations and thus the presence of threedifferent phenotypic stocks of C punctatus

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors thank the Head of the Department of ZoologyUniversity of Lucknow for providing facility and one ofthe authors is thankful to the Department of Science andTechnology (DST-PURSE) for providing him fellowship

References

[1] S X Cadrin ldquoAdvances in morphometric identification offishery stocksrdquo Reviews in Fish Biology and Fisheries vol 10 no1 pp 91ndash112 2000

[2] A Kashyap M Awasthi and M Serajuddin ldquoIntraspecificdiversity of freshwatermurrelChanna punctatusrdquo InternationalJournal of Fisheries andAquatic Studies vol 3 no 2 pp 166ndash1732015

[3] J Verma A Kashyap and M Serajuddin ldquoPhylogeny basedon truss analysis in five populations of freshwater catfishClupisoma garuardquo International Journal of Science and Researchvol 3 no 8 pp 1414ndash1418 2014

[4] A Kashyap M Awasthi and M Serajuddin ldquoGeographicmorphometric variations of freshwater murrel Channa punc-tatus from northern and eastern parts of Indiardquo Proceedingsof the National Academy of Sciences India Section B BiologicalSciences pp 1ndash7 2014

[5] L Hauser G R Carvalho and T J Pitcher ldquoMorphologicaland genetic differentiation of the African clupeid Limnothrissamiodon 34 years after its introduction to Lake Kivurdquo Journal ofFish Biology vol 47 pp 127ndash144 1995

[6] D G Bembo G R Carvalho N Cingolani E Arneri GGiannetti and T J Pitcher ldquoAllozymic and morphometricevidence for two stocks of the European anchovy Engraulisencrasicolus in Adriatic watersrdquo Marine Biology vol 126 no 3pp 529ndash538 1996

[7] C Turan ldquoA note on the examination of morphometric dif-ferentiation among fish populations the truss systemrdquo TurkishJournal of Zoology vol 23 pp 259ndash264 1999

[8] N C Ujjainia and M P S Kohli ldquoLandmark-based morpho-metric analysis for selected species of Indian major carp (Catlacatla Ham 1822)rdquo International Journal of Food Agriculture andVeterinary Science vol 1 pp 64ndash74 2011

[9] P Jayasankar P CThomasM P Paulton and J Mathew ldquoMor-phometric and genetic analyzes of Indian mackerel (Rastrelligerkanagurta) from peninsular Indiardquo Asian Fisheries Science vol17 pp 201ndash215 2004

[10] A M Sajina S K Chakraborty A K Jaiswar D GPazhayamadam and D Sudheesan ldquoStock structure analysisof Megalaspis cordyla (Linnaeus 1758) along the Indian coastbased on truss network analysisrdquo Fisheries Research vol 108 no1 pp 100ndash105 2011

[11] M A Khan K Miyan and S Khan ldquoMorphometric variationof snakehead fish Channa punctatus populations from threeIndian riversrdquo Journal of Applied Ichthyology vol 29 no 3 pp637ndash642 2013

[12] F J Rholf TpsDig Version 140 Department of Ecology andEvolution State University of New York New York NY USA2004

[13] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysisrdquo Palaeontologia Electronica vol 4 article 9 2001

[14] N G Elliott K Haskard and J A Koslow ldquoMorphometricanalysis of orange roughy (Hoplostethus atlanticus) off thecontinental slope of southern Australiardquo Journal of Fish Biologyvol 46 no 2 pp 202ndash220 1995

[15] P Jolicoeur and J EMosimann ldquoSize and shape variation in thepainted turtle A principal component analysisrdquoGrowth vol 24pp 339ndash354 1960

[16] J M Humphries F L Bookstein B Chernoff G R Smith R LElder and S G Poss ldquoMultivariate discrimination by shape in

6 International Journal of Zoology

relation to sizerdquo Systematic Biology vol 30 no 3 pp 291ndash3081981

[17] F L Bookstein ldquolsquoSize and Shapersquo a comment on semanticsrdquoSystematic Biology vol 38 no 2 pp 173ndash180 1989

[18] P Sundberg ldquoShape and size-constrained principal componentsanalysisrdquo Systematic Zoology vol 38 no 2 pp 166ndash168 1989

[19] S B Saila and J M Flowers ldquoGeographic morphometricvariation in the American lobsterrdquo Systematic Zoology vol 18no 3 pp 330ndash338 1969

[20] National Bureau of Fish Genetic Resources Annual Report ofNational Bureau of Fish Genetic Research National Bureau ofFish Genetic Resources Lucknow India 2001

[21] A G Murta ldquoMorphological variation of horse mackerel (Tra-churus trachurus) in the Iberian and North African Atlanticimplications for stock identificationrdquo ICES Journal of MarineScience vol 57 no 4 pp 1240ndash1248 2000

[22] B C Pathak J I Mir andM Serajuddin ldquoMorphometric varia-tion among barred spiny eelMacrognathus pancalus (Hamilton1822) populations from the Ganges and Brahmaputra riverbasin India by using geomorphometricsrdquo Journal of Biologyvol 3 no 1 pp 15ndash20 2013

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 6: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

6 International Journal of Zoology

relation to sizerdquo Systematic Biology vol 30 no 3 pp 291ndash3081981

[17] F L Bookstein ldquolsquoSize and Shapersquo a comment on semanticsrdquoSystematic Biology vol 38 no 2 pp 173ndash180 1989

[18] P Sundberg ldquoShape and size-constrained principal componentsanalysisrdquo Systematic Zoology vol 38 no 2 pp 166ndash168 1989

[19] S B Saila and J M Flowers ldquoGeographic morphometricvariation in the American lobsterrdquo Systematic Zoology vol 18no 3 pp 330ndash338 1969

[20] National Bureau of Fish Genetic Resources Annual Report ofNational Bureau of Fish Genetic Research National Bureau ofFish Genetic Resources Lucknow India 2001

[21] A G Murta ldquoMorphological variation of horse mackerel (Tra-churus trachurus) in the Iberian and North African Atlanticimplications for stock identificationrdquo ICES Journal of MarineScience vol 57 no 4 pp 1240ndash1248 2000

[22] B C Pathak J I Mir andM Serajuddin ldquoMorphometric varia-tion among barred spiny eelMacrognathus pancalus (Hamilton1822) populations from the Ganges and Brahmaputra riverbasin India by using geomorphometricsrdquo Journal of Biologyvol 3 no 1 pp 15ndash20 2013

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 7: Research Article Phenotypic Variation in Freshwater Murrel ...downloads.hindawi.com/journals/ijz/2016/2605404.pdf · Phenotypic Variation in Freshwater Murrel, Channa punctatus (Bloch,

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology