Utility of Caudal Fin Ray Counts in Classification of Suprageneric Categories of Indian Siluroid Fishes

BuZI. zool. SurtJ. India, 4 (2) : 221-230, 1981

UTILITY OF CAUDAL FIN RAY COUNTS IN CLASSIFICATION OF SUPRAGENERIC CATEGORIES OF INDIAN SILUROID FISHES

RANI SINGH AND K. C. JAYARAM ZoologicaZ Survey of India, Oalcutta

ABSTRAOT

The practice of routine counting of branched and unbranchcd rays of paired and unpaired fins, in fishes have been in vogue since many years. However, in recent years plotting these counts as frequency disttibution polygons have yielded a better picture in interpreting the systematic affinities and ultimately the phylogeny of the families themselves. The caudal fin in most genera of catfishes (Order Siluriformes) is forked and the number of principal unbranched rays are usually one on each lobe. An examination of a large series of material of many Indian genera and species have revealed an interesting pattern. It is seen that most families have constantly seven branched rays in the upper lobe of the caudal fin and eight in the lower lobe.

INTRODUCTION In recent years, this character has come Order Siluriformes comprises thirteen into prominence for tracing intergeneric and families, 48 genera and about 160 species from family affinities. Notably many American Ichth India, Pakistan, Bangladesh, Burma and Sri yologists have adopted this character profitably. Lanka. Most of them are confined to fresh Ginsburg (1945) was the first to demonstrate water except Ariidae and Plotosidae which the utility of different kinds of caudal fin rays occur in the sea and brackish water. Many in classical taxonomy. Gosline (1961) consi workers have studied the morphology dered only the branched rays in Cyprinoid and osteology of catfishes (Bamford, 1948; fishes and indicate that the round tailed mem Bhimachar, 1932, '33 ; Chranilov, 1929 ; David, bers have the lowest (and apparently the 1935; Eaton, 1948; Gauba, 1962, '66, '67; most variable) number of branched caudal '70; Jayaram, 1955, '70; Joseph, 1960; fin rays. Kindred, 1919; Karandikar and Masurekar, Generally, the caudal fin of most Indian 1954; Lenous, 1968; McMurrich, 1884; genera of siluroid families such as Bagndae Masurekar, 1962; Nair, 1938; Nawar, 1954; (except genus Pseudobagrus), Pangasidae, Reichel, 1927; Shelden, 1937 ; Sriniva§achar, Ariidae, Schilbeidae, An'l.blycepidae, Sisoridae 1957, '58, '59, '80 and Tilak, 1961, '63,'64, '67). (except genus Euckiloglani8, Glyptosternum, However, not much attention seems to have Ooraglani8, Oreoglanis and Pseudecheneis) and been paid towards the utility of caudal fin ray Siluridae (except Silurus) is forked. In other count as a possible taxonomic tool, families Olyrldae, Plotosidae, Chacidae Clarii- 13 III Bulletin 0/ tke Zoological Survey oj 1Mia dae, Heteropneustidae and in genera SilurU8 tions · present in the Zoological Survey of (Siluridae), Euchiloglanis, Glyptosternum, "India, Calcutta were also examined. Skeletal OoragZanis, Oreoglanis and P seudecheneis preparations of the entire caudal vertebral com.. (Sisoridae) and in Paeudobagru8 (Bagridae), plex besides Alizarine transparencies were made. the caudal fin is truncate, rounded or emargi- The different kinds of caudal fin rays enume nate. rated ~ in this study are segmented branched rays, segmented unbranched rays, and unseg MATERIAL mented unbranched rays (=simple rays). We have examined the representatives of almost all families of Indian Siluroids except The counts were made on both sides of ing Akysidae, 402 specimens falling under the caudal fin under a stereoscopic Binocular

Fig. 1. A. Caudal:fin of Auchi1WgZanis biscutatus showing 7 +8 segmented branched rays. B. Caudal fin of SiSOT rhabdophorus showing 4 + 5 segmented branched rays.

12 families 28 genera and 60 species were microscope and the data recorded. In the studied. Wherever possible fresh specimens case of unique specimens, or difficult to obtain were collected and examined. The preserved species, counts were obta,ined from X-ray specimens in the National Zoological Collec- photogravhs, SiNGH & ]AYARAM: Utility oj caudal ftn 223

RBSULTS Clariidae and Heteropneustidae 7 + 7 (fig. 2C) Table I presents the general pattern in the and Ariidae 6+ 7 (fig. 2 B) in the upper and material examined. Out of 12 families of lower half respectively. In Plotosidae and Siluroids examined six families (i.e., Amblycepi Chacidae there is no demarcation of the dae, Bagridae, Pangasidae, Schilbeidae, upper and lower half and all the caudal fin Siluridae and Sisoridae) have 15 segmen rays originate from a single hypural plate ted branched rays arranged in one pattern, (see fig. 3). As such the total number of i.e., 7 rays in upper half and 8 in the lower branched rays are 9 to 10 in Plotosidae and 10 half of caudal fin (see fig. 1 A). This frequency to 11 in Chacidae. is constant and is not seen varying even up It may be stated here that some abbera ..

Fig. 2. A. Caudal fin of Olllra longicaudata showing 5 + 6 segmented branched rays. B. Caudal fin of Tachllsuru8 dussumieri showing 6+7 segmented branched rays. C. Caudal fin of Claria3 batrachus, showing 7 +7 segmented branched rays.

to famUy level. The second group of the tions which appear to be extremely rare and families (Olyridae, Clariidae, Heteropneu isolated are also seen. Out of 26 specimens stidae and Ariidae) have 11 to 14 segmented of Olarias batrachus (Clariidae), three speci branched rays in the caudal fin ; but unlike mens (101 mm, 169 mm and 210 mm in the earlier group the number of rays in the standard length) were found to have 6 instead upper and lower half are in three different of 7 branched rays in the lower half of the patterns. Family Olyridae has 5 + 6 (fig.2A); fin. Similarly, among 16 specimens of H. 224 Bulletin, oj t}~e ZooZogical Swrvey oj I n,d,ia,

/088ilis (Heteropneustidae), one specimen (150 Considering the fact that the adipose fin is mm in SL) was found to have 6 branched also modified in the form of a spine and the rays in upper and 6 in lower half instead caudal penduncle is narrow like a whip only of 7 branched rays in both the half. The in this genus as compaTe~ to all other Indian sisorid genus Euchiloglanis generally has 7 + 8 Siluroids, the presence of only fewer segmented branched rays in the upper and the lower branched rays is significant. It would seem

0" 0 a'

Fig. 3. Caudal fin of Plotosus Zineatus with some vertebrae showing segmented branched rays.

half of the caudal fin but we have observed that Sis or represents a separate phyletic line 7 + 7 in one specimen of E, kamenganais from the other sisorids and alone perhaps (52 mm in SL). Six specimens of Eutropiichthys have to be segregated as a separate family. vacha (Schilbeidae) examined, one specimen(70 mm in SL) was found to have six segmented DISCUSSION branched rays in upper as well as lower half of The analyses of the above frequency distri caudal fin instead of 7 and 8 rays respectively. bution of caudal fin ray counts (see Table II) Genus Sisor presents some interesting indicate that the Siluroid genera examined results. The upper half of the caudal fin has by us can be grouped into five groups, based one segmented unbranched ray and four on the number of segemented branched rays. segmented branched rays; the lower half has The first group of six families, Bagridae an4 two segmented unbranched and five segmented five other mentioned earlier has 15 (7 + 8) branched rays (see fig. 1 B). The occurrance branched rays in the caudal fin except genus of nine segmented branched rays in total Sisor which comprises only 9 branched rays. in the caudal fin of this genus versus 15 Clariidae with Heteropneustidae as second in all other genera is an interesting findings. member has 14 (7 + 7) branched rays ; At'iidae, SINGH &.. JAYARAM : Utility oj caudat fin 225

TABLE-I Frequency distribution of different kinds of caudal fin rays in Indian Siluroid fishes

---...------Family, genera. and No. of Range of Total number of Total number of Total number of speoies exam- SL in mm segmen ted bran- segmented un- unsegmented pIes ched rays in branched rays unbranched upper & lower in upper and rays in upper & ha.lf lower half lower half

PLOTOSIDAE Plotosus P. Zineatus 9 92-288 9-10 P. canius 8 74-297 9-10 OHAOIDAE Chaca C. chaco, 12 81-160 10-11 *21-28

ARIIDAE Tachysuru8 T. artus 18 107.50-345 6+7 2-4+8-4 11-18+ 11-18 T. maculatus 9 112-815 6+7 8+3-4 18-19 + 11-19 T. jeZZa 4 130-225 6+7 8+8-4 13-15 + 11-18 T. gagoro, 12 198-250 6+7 2-3+3-4 11-15+ 13-17 T. thalassinus 10 116-610 6+7 2-3+3-4 15-16+12-16 T. caeZatus 18 45-190 6+'1 2-8+3-4 14-17 + 15-16 T. dussumi6rt 20 110-885 6+'1 2-4+8-4 13-16+12-16 T.SOM 6 95-325 6+7 3+3-4 11-17 + 11-16 T. sagO'l' 4: 145-250 6+7 2-8+8-4 12-15+ 12-17 T. subrostratus 10 110-285 6+7 4+4-5 11-13 + 10-12 T. crossocheiZus 5 170-245 6+7 8+8-4 17-20+17-20

Osteo~enelo8us O. militaris 14 75-865 8+'1 2+8 12-18 + 11-16 Batrachocephalus B. mino 1 223 6+ '1 4+5 11+12 OLARIIDAE Clarias O. batrachus 26 66.5-229.0 7+6··-'1 1-3+1-3 2-4+2-4 HETEROPNEUSTIDAE Beteropneustes H·fossilis 16 91-225 6··-7+6**-7 1-4+1-2 3-9+1-5 H. microps 1 98.5 7+7 4+2 4+1 BAGRIDAE Rita R. rita 15 41.5-235.0 7+8 2 .. 4+2-5 12-15+ 12-16 R. kuturnee 3 143-179 7+8 2+2-4 16-17 + 11-15

The number with * marks represents the dorsal procurrent part of ca.udal fin. 226 Bulletin of the ZoologicaZ Survey oj India

Table 1 (contd.)

Family. genera, and No. of Ra.nge of Total number of Total number of Total number of species exam- SL in mm Begmen ted bran- segmented un- unsegmen ted pIes cbed rays in branched rays unbranched upper & lower in upper and rays in upper & half lower half lower half

Horaba~rus H. brachysoma 5 96-187 7+8 2+2-8 10-15+8-14 MyatuB M. guZio 7 47-106 7+8 2+2-3 12-15+8-14 M. cavasius 5 68-108 7+8 2+2 7+7-8 Aorlchthya A.aor 5 180-198 7+8 2+2-8 8-10+6-8 SISORIDAE Glyptothorax G. pectinopterus 14 51-143 7+8 1-2+1-2 6-3+6-12 G. punjabensis 5 42-106 7+8 3+2 2+4 Glyptosternum G. reticulatum 4 57-117.5 7+8 ~+S 8+8 Euchlloa1anls E. Jeae 4: 50-110 7+8 ~+2 4-7+5-7 E. kamenganris 6 5~-115 7+7**8 2+2 5-8+5-8 E. hoago,rti 8 85-115 7+8 ~+2 5-7+5 Gaaata G. cen'ia 10 45-55 7+8 8+3-4: 10-12+10-1S G. gagata ~ 110-121 7+8 4+5 18-15+12 Nanara N. nangra 4: 46.5-75 7+8 8+4-5 11-14+18 N. 't1ridescens ~ 47-60 7+8 3-4+4-5 12-13+ 11-14 Paeudecheneis P. sulcatus 4 105-128 7+8 2-8+2-3 6-7+5-8 Baaarius B. bagarius 4 95.5-141 7+8 2+2-3 8+9-18 Sisor S. rhabdophorus 78-150 4+5 1+2 1+1

SILURIDAE Wallaao W. attu 8 135-481 7+8 1-2+1-2 ~-5+2-3 Ompok O. bimaculatus 4 115-207 7+8 1-2+1-2 2-3+1-2 O. pabo 2 106-199 7+8 1+2 2-3+2 Sllurus S. gangetica 7 75-106 7+8 2+3 14-18+14-15 SINGH & JAYAl\AM: Utility 0/ caudal lin 227

Table 1 (con td.)

Fa.mily, genem and No. of Range of Total number of Total number of Total number of species exam- SL in mm segmented bran- segmented un- unsegmen ted pIes ched rays in branched rays unbranched upper & lower in upper and rays in upper & half lower half lower half

PANGASIDAE Pangaslu8 P. pangasius 24 41-170 7+8 2-8+2-4 12-20+ 10-16 SCHILBEIDAE. Cluplsoma, O. garua 10 75-129 7+8 2-3+3-4 6-18+5-18 Eutropllchtbys E. vacha 6 70-105 6*"'-7 +6"'*-8 3+4 7-13+7-18 AUla A. coila 5 58-81 7+8 3-5+3-4 10-13+ 13-14 A. punctata 9 69-136 7+8 3-4+2-5 5-16+8-12 Sllonia S. rilonaia 8 55-80 7+8 2-8+2-4 18-17+7-13 Al\IBLYCEPIDAE Amblyceps A. mango"s 18 36-1015 7+8 8-6+8-5 9-16+6-18 OLYRIDAE Olyra O. Zongtcauaata 10 46-90 5+6 3-4+2-3 5-2+6-10

the sole member of the third group has 13 TABLE II-Frequency distribution of segmented (6+ 7) branched rays and Olyridae with 11 branched rays of caudal fin in Siluriformes (5 + 6) branched rays. The Plotosidae and 9 10 11 12 13 14 15 Chacidae form a separate group with 9-11 PLOTOSIDAE (19) br.anched rays as a single unit. CHACIDAE (12) OLYRIDAE (10) For a number of years, Regan's (1911) ARIIDAE (130) proposition that the Bagridae formed a basic CLARIIDAE (26) line of evolution of other siluroid families HETEROPNEUSTIDAE (17) has been accepted. Gosline (1944) considered BAGRIDAE (40) SISORIDAE (66) the evolution of different siluroid families not SILURIDAE (19) from a single phyletic line but from several PANGASIDAE (24) branches. Recent researches such as those SCHILBEIDAE (88) of Chardon, 1968; Srinivasacher 1980 ; show Al\1:BLYCEPIDAE (18) that there are divergent phyletic groups in N umber in parantbesis after each fa.mily indicate the Siluroids. Our studies indicate five different number of specimens examined.

----_. -----.. ------The number with •• marki represents aberra.nt number. 228 Bulletin 01 the Zoological Survey of India groupings of the families based on the fre CHRANILOV, N. S. 1929. Beitrage zur kenntnis quency distribution of the branched rays (see das Weberschen Apparates der Ostario Table II). Of these the family Bagridae appears physi. 2. Der Weber'schen Apparates to have the largest number of allied families bei Siluroidea. Z~ol. Jb. Anat., 51: compared to Plotosidae or Clariidae. It is 323-462. also evident that Plotosidae and Ariidae have separate lineages unlike Bagridae. Whether DAVID, L., 1935. Die Entwicklung der the five trends exhibited by the fin rays Clariiden und ihre verbreitung. Bine reflect the phylogeny or not is debatable but anatomisch-systematische Untersuchung. would definitely be indicative. Further work Revue Zool. Bot. air., 28 (1) : 77-147. on osteological and myological evidences are EATON, T. H. 1948. Form and function in being carried out by the authors which may the head of the channel catfish (1 ctalurU8 help to adduce definite conclusions. lacuBtris punctatus). J. ll[orph., 83: 181 .. 194. ACKNOWLEDGEMENT GAUBA, R. K. 1962. The endoskeleton of The authors are grateful to the Director, Bagarius bagarius (Ham.). Part 1. Agra Zoological Survey of India, Calcutta, for Univ. J. Res. (Sci.), 11 : 75-90. providing facilities and his sustained encoura gement. We are also thankful to Dr. William -. 1966. Studies on the osteology of Indian A. Gosline, University of Michigan for his sisorid catfishes. II. The skull of comments on the manuscript. Glyptothorax cavia. Oopeia: 802-810, 5 figs. REFERENCES -. 1967. Studies on the skull of the BAMFORD, T. W. 1948. The cranial deve Indian sisorid catfish, Erethi8te8 pU8silus. lopment of Galeichthys jelis. Proc. wol. J. Zool., Land., lSI: 379-388, 16 figs. Soc. Land., 118 (2) : 364-391. --. 1970a. The osteology of family Chaci BHlMACHAR, B. S. 1932. The siluroid skull. dae. 1. The skull of Chaca chaca (Hamil Ourf. Sci., 1 : 70-71. ton). Monitore wol. ital., 4 (1) : 21-40, 5 -, 1933. On the morphology of the skull figs. of certain Indian Catfishes. Ha11-yrly. --. 1970b. Contribution on the skull of a J. M Y80re Univ., 7 : 233-267. catfish, Erethistoides montana Hora. Zool. BRIDGE, T. W. AND HADDON, A. C., 1893. Beitr., IS: 181-201. Conttibutiontothe anatomy of Weberian ossicles in the siluroid fishes. Phil. --. 1970c. On the cranial osteology of two Trans. Roy. Sao. Lond., (B) 84 : 65-333. Indian catfishes of the genus Laguvia. Zool. Anz., 18S ; 55-67, 5 figs. CHARDON, M. 1968. Anatomie comparee de l'appareil de Weber et des structures GINSBURG,!. 1945. Contribution to a metho connexes chez les Siluriformes. M usee dology in the caudal fin ray count of Royal de L'alrigue Serie In-So-Sciences fishes and its use in classification. Oopeia, Zoologigue8-no169 : 1-125. ; 133-142. SINGH &. JAYABAM: Utility oj cauda.lJi'lt 229'

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