RESEARCH ARTICLE ISSN: 2319-6955

A SYSTEMATIC ASSESSMENT OF THREE SPECIES OF GOBIOID

FISHES (TELEOSTEI: PERCIFORMES: GOBIOIDEI) FROM KERALA

PRAVEENA M., CHANDRAN A.1, SANTHOSH S., BADUSHA M. AND YUNUS H.

P G and Research Department of Zoology, NSS College, Pandalam, Kerala, India

1Department of Fishery biology & Aquaculture, SALS, Pathanamthitta, Kerala, India

Correspondence to: [email protected]

Abstract: Analysis of selected body characters of the three gobioid species Glossogobius giuris, butis and fusca from Kerala studied revealed a clear propensity of some of the characters towards each species sufficient enough to discriminate each other. Euclidean one way single linkage cluster analysis based on the selected body characters revealed two distinct clads. E.fusca forming a single clad, whereas B. butis and G. giuris forming the second clad with high Bootstrap support (BP=100). The second clad is composed of two linkages representing B. butis and G. giuris having high bootstrap support (BP=100). The two species remains as a sister group sharing many characters. The same pattern of relationship was noted among the three species using head as well as meristic characters.

Keywords: Clad, Cluster analysis, Gobioid fishes, Meristic, Morphometrics

INTRODUCTION

Morphometrics study the co- and Gobioinellidae with variation between the patterns of sister to that clad of three families. morphological variations (Bookstein, 1991). Considering the systematics and The most common approaches referred to as of gobiids, the most notable contributions traditional morphometrics, are only a few are those of Hoese and Allen (1983), Hoese decades old (Marcus, 1990; Rohlf and and Gill (1993), Winterbottom (1993), Marcus 1993; Adams et al., 2003). The Larson (2001), Thacker (2001, 2011 a,b), family Gobiidae is the largest marine fish Cheng et al., (2005), Tornabene and Pezold family with separate spinous and rayed (2011), etc. dorsal fins and pelvic fins often joined into a In the molecular phylogeny of disc. Recent molecular phylogenetic several groups of gobioid fishes the analysis (Thacker, 2009) have demonstrated contributions of Thacker and her group that is sister to a clad containing (Thacker, 2003, 2009; Thacker and Cole,

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2002; Thacker and Hardman, 2005; Thacker divergence of characters among three and Unmack, 2005; Thacker et al., 2007, species of gobioids namely, Glossogobius 2008a, b, 2010, 2011; Thacker and Pezold, giuris, Eleotris fusca and Butis butis 2011; Thacker and Roje, 2011), Neilson and representing the families Gobiidae, Stepien (2009) and Wang et al., (2001) Eleotridae, and Butidae respectively of deserve special mention. The only recent gobioid fishes from Kerala, using land-mark monographic compilation of Indian gobioid based morphometrics, taking into fishes is that of Jayaram (1999). The main consideration 64 morphometric characters objective of the present study to evaluate the and 22 meristic characters. morphological diversity, intergradations and MATERIALS AND METHODS

Specimens of Glossogobius giuris, character with respect to standard length, Butis butis and Eleotris fusca were collected and head characters in relation to head from Kayamkulam, NTPC Bundh, length were calculated and expressed in Alappuzha. Ten specimens of each of these percentage as follows: species were examined for the present study. 퐵표푑푦 푐ℎ푎푟푎푐푡푒푟 푋100 Morphometric measurements were taken 푆푡푎푛푑푎푟푑 푙푒푛𝑔푡 ℎ 퐻푒푎푑 푐ℎ푎푟푎푐푡푒푟 using an electronic caliper. All the 푋100 퐻푒푎푑 푙푒푛𝑔푡ℎ measurements were taken following the To study the variations in morphometric and methodology of Hubbs and Lagler (1958). meristic characters among the three species Altogether 47 body characters, 22 meristic of gobiids, Principal Component Analysis characters and 17 head characters were (PCA) was performed. All the analysis were taken from each fish of all the species carried out using the statistical package studied. PAST (version 2.17c), (Hammer et al., For comparison of morphometric 2001). characters the proportion of each body RESULTS AND DISCUSSIONS

In fish systematics, the characters morphometric characters and meristic that are of prime importance are characters. Gobioid fishes belonging to the

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order of Perciformes, comprise an important Eleotridae this distance is short. In the

and most speciose group of Acanthomorpha. present study also this character, base of Larson and Murdy (2001) in their systematic second dorsal fin stands out more account of gobioid fishes of the Pacific prominently in the case of G. giuris Talwar region while relying more on meristic and Jhingran (1991) and Jayaram (1999) in characters in separating species of Gobiidae their diagnosis of Glossogobius reported a and Eleotridae, referred to one narrow isthmus and first dorsal inserted half morphometric character namely, length of or three- fourth of pectoral fin. The pectoral the base of second dorsal fin . In Gobiidae, length stands out clearly in E. fusca, according to Larson and Murdy (loc.cit.) whereas the size of isthmus, referred to by base of second dorsal fin is much longer Thalwar and Jhingran (1991) and Jayaram than the distance from the end of second (1999) probably corresponds to the character dorsal fin to base of caudal fin, whereas in lower jaw to isthmus of the present study, which recorded more prominence in E. fusca Figure 1 Principal component analysis of three species of Gobioid based on selected body characters than in G. giuris and B. butis.

C 1.6 Table 1 : Principal component ordination of

1.2 Gobioid species based on the morphometric o BPO measurement. 0.8 m B.b PL BWD FCF FAO P C-1 P C- 2 P C-3 PAS SOA p 0.4 PD Eigen 26.223 11.21 1.392 OV E.f ISA n PFL BWAI Body value 0 -4.8 -3.6 -2.4 -1.2 1.2 2.4 3.6 4.8 e PAL character -0.4 % 58.274 24.91 3.093 SO LPVL n FIL varianc 2 t -0.8 FOA e G.g -1.2 Eigen 12.51 3.40 0.66

2 -1.6 Head value character % 73.60 20.03 3.86 -2 Component 1 varianc e Eigen 9.44 5.56 9.9 Meristic value character % 62.96 37.04 1.3 varianc e Principal Component Analysis based components as most significant (Table I). on body characters of the three species of The two components together comprised gobioid fishes yielded 29 components and 83.18% of the total variance. Among the 45 eigen values, indicated the first two variable body characters studied 33 showed

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positive loadings and 12 negative loadings selected body characters of the three species in PCA. The first component having 58.27% studied revealed a clear propensity of some of total variance with the highest eigen value of the characters towards each species of 26.22 with high loadings for 17 out of 45 sufficient enough to discriminate each other. characters studied. The second component In the latter two species these characters of the showed negative component scores remained more or less comparable. B. butis, for both PC 1 and PC 2 and is placed on the on the other hand, recorded much left quarter of the ordination matrix (Fig 1). dominance in such characters as pre pectoral length, first dorsal insertion to caudal fin E. fusca recorded positive base and second dorsal insertion to caudal component scores for majority of the fin base, whereas G. giuris showed much characters in PC 1and PC2, and occupies the specificity in the character second dorsal fin right upper half of the matrix. Analysis of base. Figure 2 Principal component analysis of three species of Gobioid based on selected head characters .

Compnent 2 3.2

B.b 2.4 OW 1.6 UJL IOW PNLMHW 0.8 E.f SPO HDP SL INWHDN -2.4 -1.6 -0.8 0.8 HWNCD1.6 2.4 3.2 4 HDO

-0.8 GWPBL 2 Component 2 Component LJI -1.6 G.g -2.4POL

Component 1

Altogether 17 head characters were accounted for 73.60% of total variance with considered for the present analysis for a high eigen value of 12.5%. The second predicting the variations in head characters component explained 20.03% of the total of the species studied and recording variations in head characters with an eigen altogether 93.63% of total variation in head value of 4.40. The characters pre occipital characters. The first component alone length and orbit width were the most

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contributing variables in this component. No characters namely, prepectoral length, significant variations were noted in any of second dorsal insertion to caudal fin base the characters in the third component. As in and second dorsal fin base with high factor the case of body characters, the component loadings. The third component, on the other scores relating to head characters of each hand, recorded 3.09% of total variance individual species of fish analysis involving (eigen value 1.39) without any significant all the body characters explained 24.91% of factor loadings for any of the characters the total variance, with an eigen value of studied. 11.21. This component showed three

Figure 3 Principal component analysis of Figure 4 Euclidean single linkage one way (A) and three species of Gobioid based on meristic two way (B) cluster analysis based on the characters morphometric body characters of three gobioid fishes

G.g 2.

AL1.8 U BAR S 1. LPS P LTS E.f 0. P R ASRP - - - 0. FSLLS1. 2. 3. 4 UTR Component2 - D CPCF - TBS BTRBPR- B.b - - Component 1

The character loadings remained showed negative component scores for PC 1 very high for all the variables with an eigen and positive component scores for PC2, and value of 15.09 and 88.75% of total variance is grouped on the upper left quarter of the within the samples B. butis, for instance, component matrix. G. giuris, on the other

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hand, obtained for PC1 and PC2 are half of the ordination matrix. Analysis of represented in a two dimensional matrix in head characters clearly indicate an obvious Fig 2. B. butis, for instance, with negative segregation of characters specific to each component scores for PC 1 and positive species of fish studied and majority of the component scores for PC 2 is located on head characters remained more pronounced upper left quarter of the component matrix, in E. fusca than in the other two species, and whereas G. giuris with negative component seemed to be so characteristic of the species. scores for both PC 1 and PC 2 is located The species B. butis, on the other hand, opposite to B. butis on left lower half of the showed dominance in characters such as component matrix. In the case of E. fusca snout to preopercle, snout length and orbit both the components PC 1 and PC 2 had width, whereas the character preoccipital positive scores and this species is located length recorded more specificity in G. giuris away from the other two on the middle right than in E. fusca and B. butis.

Figure 5 Euclidean single linkage one way (A) and two way (B) cluster analysis based on the morphometric head characters of three gobioid fishes

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Principal Component Analysis was pelvic scale rows and pre anal scales, is performed using the 15 meristic characters. plotted opposite to B. butis on the upper left The first component explained 62.99% of quarter of the scatter plot (Fig. 3). According total variance with an eigen value of 9.44 is to Thacker (2009) the newly erected family highly significant. The second component Butidae are sister to Gobionellidae plus recorded the rest of the total variance (37.04 Gobiidae, with Eleotridae forming another %) of the characters with an eigen value of major clade, and Rhyacichthyidae and 5.56. In this component only three variables Odontobutidae remaining sister to the namely, branched anal fin rays, branched remainder of Gobioidei. pelvic fin rays and pre anal scales, Figure 6 Euclidean single linkage one way contributed most with high loadings. The (A) and two way (B) cluster analysis based species, E. fusca with highest values for on meristic characters of three gobioid fishes many of the variables is placed in the right half of the scatter plot. All the variables so characteristic of E. fusca had high positive loadings in both PC 1 and PC 2. The species Butis butis, on the other hand, with high values for unbranched pectoral fin and pre anal scales and lowest value for pelvic fin spine, branched pelvic fin rays, lateral line scales, pre dorsal scales, upper transverse scale rows, circumpeduncular scale, circumferential scales and transverse breast rows is plotted on the lower left quarter of the ordination matrix. G. giuris with highest values for branched pelvic fin rays and branched pectoral fin rays and lowest values for branched anal fin rays, lateral line to In the present study, Euclidean one selected body, head and meristic characters way single linkage cluster analysis based on of the three species of gobiids, yielded two

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distant clads; E. fusca forming a single clad is highly congruent with that reported by whereas G. giuris remains sister to B. butis Thacker (2009) for Gobioid fishes. together forming the second clad with high Humphries et al., (1981) pointed out that a Bootstrap support (BP=100). The second bridge is needed between traditional clad is composed of two linkages quantitative methods and geometric analysis representing B. butis and G. giuris having of shape. Mekkawy (2002), on the contrary, high bootstrap support (BP=100). The two opined that traditional characters are more species remains as a sister group sharing reliable than the truss ones for some species. many characters (Fig. 4, 5 and 6). This During the present study both the traditional relationship, in terms of the respective as well as the truss characters were taken in families, Eleotridae, Gobiidae and Butidae, to consideration.

CONCLUSION

The species selected for the present forming a single clad whereas B. butis and study were Glossogobius giuris, Eleotris G. giuris remaining as a sister group sharing fusca and Butis butis. Altogether 47 body many characters. The results of the present characters, 17 head characters and 22 study are compared and discussed at length meristic characters were taken from each with those of similar studies available for fish. All the data were subjected to Principal gobioid fishes from other geographical Component Analysis which yielded areas. Detailed morphological and characters that could be used for separating phylogeographic studies involving more the three species of gobiids. Cluster analysis genera and species of gobiids using both employing morphological (body and head) traditional as well as modern taxonomic characters, as well as meristic characters tools are suggested. revealed two distinct clades, E. fusca

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