Arius (Pisces : Siluriformes : Arfidae)
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Proc, Indian Acad. SCi. (Anim. Sci.), Vol. 91, Number l, January 1982, pp. 79-98. ~ Printed in India. Some biometric studies of certain closely related species of the genus Arius (Pisces : Siluriformes : Arfidae) J R DHANZE and K C JAYARAM Zoological Survey of India, 27 J L Nehru Road, Calcutta 700 016, India MS received 6 March 1981 ; revised 17 August 1981 Abstract. The marine catfish genus Arius of the family Ariidae comprisrng 21 species have been divided into six complexes and three groups based on interspecific relationships and morphometric affinities. In this paper the maculatus complex of four species, viz., Arius maculatus, Arius arius, Arius gagora and Arius jella has been critically examined in respect of a selected list of 20 morphological characters based on examination of a large series of examples collected first hand bY the authors. The samples have been statistically analysed, and the range of variation in respect of each character as exhibited by each species has been delineated. The probability significance test has been made to establish the interspecific relationship. Keywords. Biometric study; Arius species; Ariidae, 1. Introd1lCtioQ The genus Arius Valenciennes, 1840 forms a commercially important group of marine catfishes comprising 21 species from India, Pakistan, Bangladesh, Burma and Sri Lanka. Most of the species are marine often entering estuarine waters and occasionally even in freshwaters such as A. acutirostris, A. burmanicus and A. gagora etc. About 80% of the total catfish landing in our country is of Arius species. Despite the economic value of these fishes, the taxonomic identity of most of the species is in a state of confusion. The main reason for such ambi guity is because earlier workers depended mainly on one or two characters which were highly variable interspecifically if not associated with the changes in growth or sex. Day (1877, 1889) gave a comprehensive account of 23 species by using the anal fin counts, relative head length and eye diameter as diagnostic characters, besides the shape and size of teeth bands on the palate. Weber and de Beaufort (1913) also utilised the dentition pattern, besides the shape of the occipital process for separating the species of this genus. Smith (1945)considered the dentition pattern as one ofthe very important taxonomic character and stated, "the most important character for separating the species are teeth." Chandy (1954) framed a key mainly based on the dentition pattern on the palate, for the identification of Arius species present in the NZC of ZSI, Calcutta. Subsequent ichthyologists also relied upon this character (Munro, 1955 ; Smith, 1962; Wongratana and Bathia, 1974 ; Misra, 1976). Taylor (1978) adopted the length of the median 79 80 J R Dhanze and K C Juyarani longitudinal groove on the head, the shape of the bony shield for separating Arius species of western central Atlantic (Fishing Area 31). It may b e seen that for separating the various species of Arius the pattern of teeth patches on the palate still remains to be an unavoidable necessity. How ever, it may be indicated that whereas the basic contour, the number and posi tion of the patches remain constant, the size, number and nature of the teeth themselves vary highly and alter considerably with age and growth. Earlier ichthyologists seem to be unaware of this fact and established species like A. serratus Day, A. malabaricus Day, A. satparanus Chaudhuri for such variants which are invalid (Jayaram and Dhanze 1978a~ 1981). Based on the number and contour of the patches we have placed the 21 species of Arius in six complexes under three groups. The constituent species of each complex are closely interrelated and some may even prow later either to be synonyms or subspecies. In this paper 'maculatus' complex which has four species (A. maculatus, A. arius, A. gagora and A. jella) (Text-figures 1A~H), have been analysed to determine their interspecific affinities and systematic status. 2. Materials and methods The material for this study is based on 430 specimens collected by the authors during extensive survey tours of the entire eastern. coast and ~ part of southwest 1: A \'. -\' '·:;'i:::·:;·:;;::::!:'i;:~~f:i·l· H Text Figure 1 A.-A. maculatus (dorsal view of head). B.-A. maculatus (dentition). C.-A. gagora (dorsal view of hcadj .; D.-A. gagora (dentition). E.-A. arius (dorsal view of hoad). F.-A. arius (dentition), G.-A. [ella (dorsal view). H. A. jela (dzntition). (Figures A-E, G and H, after Chandy, 1953, Figure F-after Chaudhuri 1916). Biometric studies of the genus Arius 81 coast of India. The specimens presen t in the National Zoological Collections of the Zoological Survey of India, Calcutta, have also been examined. Fresh material of species SUC]1 as A. jella and A. maculatus were collected and studied by the second author (KC1) during the FAO consultation, Cochin in 1980. A total of 45 characters were mensurated and of which 20 alone are selected for the statistical analysis. All the measurements were taken with dial calipers to the nearesthalfof a millimeter for the size range upto 150 mm and by measuring tape abcve this size. Taxonomic characters are generally found to intergrade or overlap between closely related species when a large series of specimens are studied. The reliability or otherwise of such characters are to be evaluated. Different methods of measuring intergradation 01' divergence have been proposed (Davenport and Blankinship, 1898; Pearl, 1930; Ginsburg, 1938 ; Simpson and Roe, 1939 ; Amadon, 1949 ; Snedecor, 1956; and Simpson et at 1960). Methods deviced tby Simpson et al (op. ca.) for the comparison of two populations irr-espective of heir taxonomic identity seems to be useful here. The "Student's z-test " to determine the probability value at 95~ confidence intervals have been applied. Before deducing any ilUm<'lical conclusion, a hypothesis was set forth that all the specimens of different populations examined belong to a same species, and the univers .lly used rejection value of 5 per cent was chosen as a criteria for the rejection of tris hypothesis. However, the estabIisf-ment of the significance of a difference between two species by numerical derivation is not in itselfa zoological conclusion. Thus the numerical expressions for each character were further compared or rather standaridized by employing geometrical expressions proposed by Dice and Leraas (1936), and later on adopted with some modification by Hubbs and Perlmutter (1942), Pillay (1951), Hubbs (1952), and WiEterbottom (1980). In this method, for each character the range, mean, one standard deviation and one standard error on each side of the mean were delineated on the graph. The degree of overlap or sepal', tion of the standard deviations in respect of the arithmetic mean of each species Was determined. 3. Results Tables 1-6 and graphs 1-20 present the biometric comparison of the four species with each other for ell the 20 characters selected. 3. 1. A. maculatus vs. A. arius It is seen that excepting the head length, in respect of all other characters the two species have a probability of less than 0 '1% and are significantly different (table 1). From the graphs XIV-XVI, XVIII, XX, Dice diagram A and Bin each, it is seen that the mean of each population as well as standard deviation (S) diverge to a considerable degree, thereby justifying the separate specific status of A. maculatus and A. arius. Both the species have a single large oval patch of teeth on each side of the palate (tex t-figure 1 B, F). Further, the two species can be morpho logics lly distinguished by the size and position of the eye The eye diameter is 1&-50% ill Ai maculatus vs. 21·40% in A. arius in the hear: length; 33·30% lS 45·60% i.i the interorbital width and 57·35% vs. 63·30% in snout length, QC Table 1. Biometrical comparison of various morphometric data for A. macula/us vs, A, arius from different localities of east and west IV coast of India, 67 s",Jccimens of A_ macula/us (Thunberg) 81 specimens of A. arius (Hamilton) ------------------ <...., Range Mean X1±SX 8, Range Mean x,±sx S. t P ::tl LHjTL % 18' 60-25' 00 21-1238±'23 1,7656 18'87-24-32 20-7239± -18 1 '3642 1'3796 10--20% ~b ,;:, HBjTL % 15'13-21'58 18' 3448±' 28 1- 9319 13-52-21'50 16-3244± '19 J '4744 6-4097 <0'1% ~ 26' 7961 ± -28 2-0935 24,65-·29'il8 26'6695± -18 1'3996 <60·70% LHjSL % 23-29-30-12 0'4185 '",." HE/SL % 18'75-26'97 n 9738± -28 2'0858 17'77-26'23 21' 0027 ± -21 1'6385 5'7282 <0')% ::> 7280 )'00% :::, PDL/SL ~~ 32-41-39-44 36'3168±'Z3 1- 35-2$-40'45 37'0227± '20 I' J529 - 2-6251 "---. PALj5L % 66,67-75'00 (i9'6174±-~1 2-)284 63-93-73-08 69'8702± '29 2'278 - 0-6025 <0-1% ='\ PPL/SL % 21 '72-30'~S ;~5'9972::!: '36 2"7055 22-40-27'41 24'707&± '1& 1'3965 - 3'28J7 0'1% ') 4,1754 61' 6339± - 3'9983 - 2-5670 <1'00% WDFjWAF % 50'00-/2'22 59'7017±'55 52' 59-69' 70 51 0..... WH/LH % 63'16-84' 09 n'8778±'6,1 ~, 8220 62,60-78' 57 70,1412± '45 3' 51167 3-5376 <0-1% 2 § HH/LH ~~ 55' 26-71' 59 65' 4108± -57 4'2906 52'00-70'00 60' 6015±1' 26 4-7358 5'4483 <0'1% LSjLH % 28: OO~37' 25 32' 3725±'29 2-1.60J 30'00-37'00 33-8687± '21 1-6373 - 4'2560 <0'1% § BDjLH % 14' 7l-22' 22 13'4687±-21 J-5800 16'00-26'67 21' 3634± ' 29 2' 2268 - 7-9n6 <0'1% INwjLH ~~ 17-91-23'86 2.1-2597±'20 1-5291 15' 29-21-77 lS'2822± '18 \ -4063 11'0167 -.o1'/'0 IOWjLH % ,H' 19-67' 39 56'1914±-67 4'5746 41'07-52:75 47'0943± '43 2-7251 12-8595 <0'\ 'io BDjLS % 39'47-66'67 57'3725±'82 6' 2239 43, 24-77 -78 63'3336±J '07 8'3560 4'3693 <0'\% INW/LS % 54-55-76- 67 {'5'9427±-aO