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A review of the genus Caranx from the tropical east Pacific Lane, E. David 1962

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A REVIEW OF THE GENUS CARANX FROM THE TROPICAL EAST PACIFIC by E. DAVID LANE B.Sc, University of British Columbia, 1959 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of Zoology We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA April, 1962 In presenting this thesis in p a r t i a l fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make i t freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It i s understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of ZOOLOGY  The University of British Columbia, Vancouver 8, Canada. Date APRIL 30, 1962 - i -ABSTRACT Seven Caranx species occur in the tropical east Pacific, _C. caballus (61 specimens examined) is separable from the Atlantic form C. crysos (10 specimens examined) by a character index. C_. marginatus (64 specimens examined) is separable from the Atlantic form C. latus (5 specimens examined) and the Indo-Pacific form C, sexfasciatus (2 specimens examined) by vertebral count. C, hippos (58 specimens examined) occurs on both sides of the Panama Isthmus. C. lugubris (28 specimens examined) is circumtropical. C_. melampygus (37 specimens examined) is synonymous with (3. stellatus. C!. medusicola (3 specimens examined) is endemic to the east Pacific region, as is C!. vine tus (22 specimens examined). Caranx species in the tropical east Pacific comprise: 2 circumtropical species (C_. hippos and C_. lugubris); 1 Pan-Pacific species (C. melampygus) and 4 endemic species (C. caballus, C. marginatus. C, vinetus and C. medusicola). Barriers to Caranx distribution in the east Pacific are: temperature to the north and south; the Panama Isthmus to the east; and the east Pacific sea barrier to the west. Some forms vary to the extent of specific difference over their range (C. caballus : _C. crysos and _C. marginatus : C. latus : C. sexfasciatus) while others are very similar over their entire range (C. lugubris and C_. melampygus). C. hippos is intermediate in variability between these extremes. Probable reasons for paucity of the tropical east Pacific fauna are the presence of barriers to invasion, especially the east Pacific sea barrier and the lack of coral reef habitat. On the basis of Caranx evidence i t is unlikely that the postulated 3 to 4C° Pleistocene temperature drop had a serious effect on the fish fauna. ACKNOWLEDGEMENT The author wishes to express his sincere gratitude to Dr. J.C. Briggs and Dr. C.C. Lindsey for suggesting and supervising this study. Also, gratitude is expressed to Dr. J.F. Bendell, Dr. B.M, Bary and Dr. N.J. Wilimovsky for their c r i t i c a l advice. The author is indebted to Mr. F.H. Berry of the U.S. Fish & Wildlife Service Biological Laboratory, LaJolla, for his encouragement and advice and Mr. J.D. McPhail for the translation of Lindberg's work on marine water levels. The specimens examined were collected by Dr. H.R, MacMillan, Dr. P.A. Larkin, Dr. C.C. Lindsey, Dr. J.C, Briggs, Dr. M.A. Newman, Mr. R.J. Krejsa, Mr. F.T, Pletcher and Mr. R.E. Johannes. Further specimens were supplied by Dr. C.R. Robins, University of Miami and Mr. W. Baldwin, University of California at Los Angeles. These people are gratefully acknowledged. - i i i -TABLE OF CONTENTS Page ABSTRACT i ACKNOWLEDGEMEM: i i TABLE OF CONTENTS i i i LIST OF TABLES v i LIST OF FIGURES v i i INTRODUCTION 1 MATERIALS AND METHODS 2 MATERIALS 2 COUNTS AND MEASUREMENTS 2 TREATMENT OF PROPORTIONAL MEASUREMENTS 4 ABBREVIATIONS 5 ILLUSTRATIONS 5 BIBLIOGRAPHIC METHODS 6 FAMILY CARANG3HAE 7 DESCRIPTION OP THE FAMILY CARANGIDAE 7 KEY TO THE GENERA 9 THE GENUS CARANX Lacepede 12 DESCRIPTION OP THE GENUS CARANX 12 DESCRIPTIONS OF CARANX SPECIES 15 CARANX CABALLUS Gunther 15 Synonomy 15 Material Examined 15 Description 16 Distribution 17 CARANX VINCTUS Jordan and Gilbert 17 Synonomy 17 Material Examined 18 Description 18 Distribution 19 CARANX HIPPOS Linnaeus 20 Synonomy 20 Material Examined 20 Description 21 Distribution 22-CARANX MARGINATUS G i l l 23 Synonomy 23 Material Examined 24 Description 24 Distribution 25 - iv -Page CARANX LUGUBRIS Poey 26 Synonomy 26 Material Examined 26 Description 27 Distribution 28 CARANX MELAMPYGUS Cuvier 28 Synonomy 28 Material Examined 29 Description 29 Distribution 31 CARANX MBDUSICOLA Jordan and Starks 32 Synonomy 32 Material Examined 32 Description 32 Distribution 33 CARANX Sp 34 COMPARATIVE MATERIAL FROM THE ATLANTIC 35 CARANX LATUS Agassiz 35 Material Examined 35 Description 35 CARANX CRYSQS Mitchell 35 Material Examined 35 Description . . . . . 36 KEY TO THE SPECIES OF CARANX IN THE TROPICAL EAST PACIFIC . 37 DISCUSSION 39 DEFINITION OF THE TROPICAL EAST PACIFIC 39 SPECIES AND SPECIES COMPLEXES 39 Caranx caballus : crysos : fusus 39 Caranx vinetus 41 Caranx hippos 41 Caranx marginatus : latus : sexfasciatus . . . . 42 Caranx lugubris 44 Caranx melampygus = stellatus 44 Caranx medusicola and melampygus 45 BARRIERS TO DISTRIBUTION 46 Panama Isthmus 46 East Pacific Barrier 47 Temperature Barriers 48 CARANX; ZOOGEOGRAPHY AND EVOLUTION 50 - V -Page SUMMARY 54 BIBLIOGRAPHY 56 APPENDICES 63 - v i -LIST OP TABLES Page I List of some characters used and the number of specimens examined 3 II Differences in some characters between (2. crysos and C. caballus 40 III Differences in some characters between C. latus and C_. marginatus 43 IV Number of spots and size of C_. melampygus 44 V Distribution and differentiation of non-endemic Caranx species in the east Pacific 51 - v i i -LIST OF FIGURES Page 1. Measurements made on Caranx species 2a 2. X-ray photograph of C. hippos 3a v 3. Length/head length and head leDgth/upper jaw length . . 4a 4. Length/depth 4b 5. Head length/eye length and length/pectoral f i n length . 4c 6. Caranx caballus 16a 7. Caranx caballus distribution . . . . . 16b 8. Caranx vine tus 19a 9. Caranx vinctus distribution 19b 10. Caranx hippos 22a 11. Caranx hippos distribution 22b 12. - Caranx marginatus 25a 13. Caranx marginatus distribution . . 25b 14. Caranx lugubris 27a 15. Caranx lugubris distribution 27b 16. Caranx melampygus 31a 17. Caranx melampygus distribution 31b 18. Caranx medusicola 33a 19. Caranx medusicola distribution 33b 20. Character index separating C. crysos and C_. caballus . 40a 21. Some proportion characters for C. crysos. C_. caballus. C. latus and C. marginatus 40b 22. Anterior profile of C. medusicola and C. melampygus . 45 23. Equatorial Pacific Ocean 47a 24. Present distribution of tropical waters (From Briggs, 1961) 48a 25. Distribution of tropical waters with a winter temper-ature drop of 3 C° (From Briggs, 1961) 48a 26. Distribution of tropical water with a winter temper-ature drop of 8 C° (Based on Sverdrup et a l , 1942) . -. 48b I N T R O D U C T I O N The following four problems are treated in this study. To attempt to determine the number of species of Caranx in the tropical east Pacific area and what these species should be named* The species pairs C. crysos and C_. caballus. C. latus and £. marginatus, C. melampygus and C. medusicola and C. melampygus and C. stellatus are of special interest here. To indicate the distribution of the specie's that are present in the tropical east Pacific both on a world wide basis and within the tropical east Pacific. To devise effective keys to the genera of the family Carangidae and the species of the genus Caranx that are found in the tropical east Pacific. To discuss the two main theories as to the reasons for the poor shorefish fauna of the tropical east Pacific and to attempt to evaluate them in terms of the genus Caranx. This entails four main points: the effectiveness of the Panama Isthmus barrier, the effectiveness of the East Pacific barrier, the emergence of the isthmus, and the effect of Pleistocene glaciation on the tropical east Pacific. M A T E R I A L S AN D M E T H O D S MATERIALS Most specimens examined came from the M.V. Marijean collections made in the tropical east Pacific between 1954 and 1961 inclusive. Some specimens were collected by F.T. Pletcher in Mexico (1959) and R.E. Johannes in Panama (1959). Fish from the Gulf of California came from the Fish Museum at the University of California at Los Angeles and comparative material from the Atlantic and Caribbean came from the Museum of the University of Miami Marine Laboratory. A tabular presentation of the material studied, size ranges and distribution i s given at the start of each species description. COUNTS AND MEASUREMENTS The fish were measured with a stainless steel Helias M.B. 1 calipers (200 mm.) calibrated to one tenth of a millimeter. Measurements exceeding 200 mm. were made on a standard measuring board using a Leitz plastic rule calibrated to 1 mm. A l l counts and measurements are as in Hubbs and Lagler (1949) except as noted below (see Fig. l ) . Fork length was used in preference to standard length due to the difficulty in establishing the posterior end of the body as there are in this region considerable adipose deposits and overlapping scutes. Caudal f i n ray counts were divided into major rays (those which reach the trailing edge of the fin) and dorsal and ventral minor rays (those which do not reach the trailing edge) (Fig. 2). G i l l rakers were counted on the lower portion of the f i r s t right hand g i l l arch, including a l l rudiments which were sufficiently developed to be f e l t with a fine probe. Lateral scutes counts include those specialized scales on the posterior half of the lateral line (straight portion), starting at the f i r s t scale with a pointed posterior 4 3 1- Fork length 4 - H e a d length 2- Depth 5 - E y e length -3 - Pectora l fin length 6 - Upper jaw length F i g u r e 1. Measurements made on Caranx s p e c i e s . edge and raised center and ending at the last scale on the lateral line which is pointed . One or two rounded scales may follow this* Spines are defined as unbranched and clear, while rays are usually branched and always cross-striated. The classification "unbranched rays" (Shults et a l , 1953) will not be used here. By these definitions, in Caranx there is one short spine on the dorsal edge of the pectoral f i n . A l l counts, measurements and characters used are based on an examination of a l l specimens listed (unless the structure involved was missing or mutilated) except as shown in Table I. TABLE I List of Some Characters Used and the Number of Specimens Examined. Character No* of Specimens of each Species Examined Relationship of ventral f i n to anus 10 Caudal major and minor rays 10 Relationship of longest dorsal spine to longest dorsal ray 10 Relationship of arched to straight portions of lateral line 10 Relationship of longest dorsal ray to the base of soft dorsal 5 Length of scaly sheath along soft dorsal and anal base 5 Osteological characters 10-40 The bone structure was examined from either prepared skeletal mounts or X-ray. The film used was Ilford Ilfex X-ray film 836X-EX744} the machine used was a General Electric Model D movable unit. Osteology varies l i t t l e within the genus Caranx. Fig. 2 is an X-ray photograph of C. hippos. Abdominal vertebrae (those in front of the anastomosed - 3a -F i g . 2 X - R a y p h o t o g r a p h of C . h i p p o s 1. D o r s a l p t e r y g i o p h o r e s p i n e 2. F i r s t a b d o m i n a l v e r t e b r a 3. F i r s t c a u d a l v e r t e b r a 4 . L a s t c a u d a l v e r t e b r a 5. F i r s t m a j o r c a u d a l r a y 5'. L a s t m a j o r c a u d a l r a y - 4 -f i r s t haemal spine and anal support) are 10 in number* Caudal vertebrae (those posterior to the abdominal vertebrae) are 14 or 15 in number (Pig. 2). TREATMENT OF PROPORTIONAL MEASUREMENTS To evaluate the usefulness of proportional characters two standard deviations were plotted on either side of the mean, using the formula (Walker and Lev, 1953). Figures 3, 4 and 5 show the ranges, 2 standard deviations (equaling 95$ confidence intervals (Snedecor, 1956)), and the means of the characters used for a l l species. Between any two species i f no overlap occurs in the plot of two standard deviations, the character plotted is valid to separate 95$ of the specimens in these species. The graphs in Figs. 3, 4, and 5 show the specimens of any given species divided into those smaller than 250 mm. and those 250 mm. and larger. From these figures i t can be seen that fork length/depth, fork length/head length and head length/upper jaw length are useful proportions to separate some species. Fork length/pectoral f i n length or any proportion involving pectoral f i n length was found to be of l i t t l e value as the very long, narrow trailing end of the f i n is subject to breakage and i t is often impossible to ascertain i f breakage has occurred. Another frequently used character which was found invalid in separating the species is head length/eye length, as was any proportional measure involving eye length. These characters are unsatisfactory as adipose tissue often make eye length d i f f i c u l t to measure. Also, the ratio between eye length and head length, or any other body measure tested using eye length varied greatly with growth. One could divide the fish into small size groups and find significance but this is not necessary in separating the species dealt with here. 2.5 3.0 3.5 J i Figure 3. Length/head length and head length/upper Jaw length, a* 2 standard deviations about ®ean (J) • d = C. marginatus 86-243 ©a 64 specimens e =* CT lugubris T97-220 so 3 specimens e'* C7 luiiubria 288-755 mm 25 specimens i i « range a a G. caballus 167-241 mm 31 specimens a 'a c . caballus 250-403 mm 30 specimens b n 57 yinctua 60-284 mm 22 specicseno c a C. hippos 66-246 m 46 specimens c' = C. hippos 262-730 mm 12 specisens f a C« aelampygua 89-241 on 29 specimens f ' n raelaapymia 398-675 as 8 specimens g = C. aedualcola 112-157 ®m 3 epecioens - 4b -F i g u r e 4. L e n g t h / d e p t h , aymbola ae i n F i g u r e 5* 2.0 2.5 3.0 — i — 3.5 — i — 4.0 4.5 5.0 — i — 5.5 6.0 _H E -1 .93 • 7 0 S -P Figure 5. Head length/eye length and length/pectoral f i n length symbols as Figure 3 except a = caballus ( a l l specimens) b = £i vinctus ( a l l specimens) c = 9JL hippos ( a l l specimens) d = marginatus ( a l l specimens) e = lugubris ( a l l specimens) f = C_j_ melampygus ( a l l specimens) g = C. medusicola ( a l l specimens) i n - 5 -ABBREVIATIONS Abbreviations used in this work are as follows: •jj = Length divided by depth •j| = Length divided by head length •j; = Length divided by pectoral length JJ = Head length divided by upper jaw length JJ — = Head length divined by eye length T.E.P. = Tropical East Pacific (3. = in front of species name equals Caranx Spec* = specimen(s) as opposed to Sp* = species x = average I. = Island; I s . = Islands B.C. Catalogue No. = from University of British Columbia W Catalogue No. = from University of California at Los Angeles U.M.M.L. Catalogue No. = from University of Miami Marine Laboratory ILLUSTRATION The maps of the Tropical East Pacific were based on those in Bartholomew (1956) reduced by squares to the appropriate size. The fish illustrations were drawn from particular specimens indicated on the drawings. The fish was placed in a tray with an elastic band around the long axis of the tray. This band was lined up with the straight portion of the lateral line and pinned to the fish thus giving a base line from which to plot points on the illustration. Where reduction in scale was necessary, points were plotted on the drawing with Dietzgen 1% inch proportional dividers. Stippling technique is that described in Cannon (1936)* - 6 -BIBLIOGRAPHIC METHODS Dean's Bibliography of Pishes, Volumes I, II, and III was examined to obtain the references from Linnaeus to 1915* The Zoological Record and Biological Abstracts were consulted from 1915 to the present* In searching through Dean many t i t l e s of the following nature were encountered, "A Collection of Marine Pish from the Pacific Coast of Mexico with some Descriptions". A l l such t i t l e s were checked, many of which made no reference to Caranx. or perhaps only a distribution record. These tit l e s are not included in the present bibliography. The ti t l e s listed in the bibliography are those cited in the text and those listed in the species synonymies. Papers listed in the synonymies, but not seen by the author, are indicated. Included in the synonymies are the original description, papers recording the geographical range of the species, and a l l descriptive and figurative references from the tropical east Pacific for the species concerned. FAMILY GARANG3DAE Caranx is a genus of circumtropical fishes of the family Carangidae, Order Percomorphi. The family was f i r s t placed in this order by Regan (1909), contested by Starks (1909, 1911), who claimed this Family should be in Order Scombroidea. Recent authors have followed Regan, Notable among such authors are Berg (1947) Perciformes = Percomorphi; Matsubara (1955) Percida = Percomorphi; and Herald (1961) Percomorphi, A family description i s included here as the existing descriptions are either too short and lacking in important details (Jordan and Evermann, 1896; Meek and Hildebrand, 1925, taken from Jordan and Evermann; Walford, 1937) or too long and unweildy (Ginsburg, 1952), The following description conforms in general to that of Roxas and Agco (1941), including the salient diagnostic features of the family. DESCRIPTION OF THE FAMILY CARANGIDAE Body more or less compressed, oblong or elongate, or short and deep. Caudal peduncle generally slender. Short ridges on the peduncle and caudal f i n above and below the lateral line in many genera. (These ridges present on Caranx and are the reason for using fork length rather than standard length)• Lateral line complete, arched anteriorly and straight posteriorly. Lateral scutes either arming the lateral line for its entire length, or only the posterior straight portion (Caranx). or entirely absent. Mouth moderate, maxillary not reaching past the eye except in Oligoplites, premaxillary usually protractile, maxillary with or without supplemental bone. Dentition varied, complete or incomplete, sometimes teeth deciduous or overgrown with flesh in larger specimens. Adipose eye shields often well developed. G i l l opening wide, g i l l membranes usually not united, free from isthmus. Branchiostegals 7. - 8 -G i l l arches 4, a s l i t behind the last. Scales complete or incomplete, cycloid, often deciduous. Dorsal fins two, the f i r s t spinous with 3-8 spines, moderate in height to very low. The pterygiophore bearing the f i r s t dorsal spine is pointed forward giving the appearance of a procumbent spine, the term applied to this structure by other authors (See Pig. 2). This point is visible in young, grown over with skin in older specimens. Spinous dorsal usually depressable into groove. Second dorsal long and soft with 1 spine followed by 14-40 rays. Anal f i n with 1 spine followed by 14-30 rays and preceeded by two individual spines. Detached finlets occasionally present after dorsal and anal fins. Pectoral f i n short or long, where long falcate. Ventral fins thoracic, 1 spine and 5 rays. Swim bladder present. Pyloric ceaca usually extremely numerous (more than 100 in some forms). Members of this family inhabit a l l warm seas in the tropical faunal regions. In the tropical east Pacific there are 16 genera and 33 species (See Appendix l ) . - 9 -KEY TO THE GENERA Existing keys to the genera are d i f f i c u l t to use. Jordan and Everman (1896) and Meek and Hildebrand (1925) place considerable emphasis on dentition in their keys. Teeth, however, in large specimens of some genera of Carangidae become deciduous or overgrown with flesh. Walford (1937), contains some errors in counts and is unreliable in these sections. For these reasons i t was deemed necessary to devise the following new key. A Ventral outline more strongly curved than dorsal. Pectoral fins almost twice as long as head. Chloroscombrus AA Dorsal outline equally or more strongly curved than ventral. Pectoral fins shorter or longer than head. B Lateral line with bony scutes. C Dorsal and anal fins each with detached f i n l e t . Decapterus CC Dorsal and anal fins without finlets D Shoulder girdle with a deep furrow near its junction with the isthmus, a fleshy projection above i t Selar Gill C o v e r — F l e s h y P r o j e c t i o n DD Shoulder girdle without furrow or fleshy projection. - 10 -E Lateral line with bony scutes along i t s entire length. Trachurus EE Lateral line with bony scutes on the posterior straight portion only. E Dorsal and anal fins each with more than one long filament. Alectis FF Dorsal and anal fins with one or no long filament. G Bony scutes on lateral line more than twenty. H Soft dorsal with 29 or more rays. Carangoides HH Soft dorsal with fewer than 29 rays. I Dorsal and anal fins never falcate, maxillary very narrow, its greatest width scarcely ? eye. Hemicaranx II Dorsal and anal fins falcate except in C. vinetus; maxillary broad, greater than l/3 dia of eye. J Dorsal and anal fins each produced into one filament. Citula JJ Dorsal and anal fins not produced into filament. Caranx GG Bony scutes on lateral line fewer than twenty. £ Dorsal and anal fins falcate, anterior profile not near vertical. Gnathanodon KK Dorsal and anal fins not falcate, anterior profile near vertical. (The scutes in this genera are often very weak and di f f i c u l t to see). Vomer BB Lateral line without bony scutes. L Second dorsal and anal about equal in length, both longer than distance from back of head to anus. 11 M Body deep, greatly compressed; pectoral fins 1.8 times or more in head-*-; preorbitals extremely deep; snout steep. MM Body not greatly compressed. Pectorals always less than 1.5 times head^; preorbitals not deep. N Maxillaries large - never more than 2 l/3 in head; dorsal and anal fins never falcate. Scalesdeeply embedded, skin leather-like. Hilary not large - never less than 2 2/3 i head ; dorsal and anal fins falcate. NN Max m inScales not deeply embedded, skin not leather-like. Selene Oligoplites Trachinotus LL Anal f i n much shorter than second dorsal, i t s base shorter than abdomen. 0 Dorsal and anal each with a single detached f i n l e t . Elagatis 00 Dorsal and anal without finlets. P First dorsal with 6-8 slender spines. Seriola PP First dorsal with 3-4 low s t i f f spines. Naucrates This character is uncertain, but the pectoral f i n of Selene is much longer than that of either Oligoplites or Trachinotus. This character i s also somewhat uncertain for some species, but the maxillary is much larger in Oligoplites than in Trachinotus. THE GENUS CARANX Lacepede The genus Caranx was f i r s t designated by Lacepede (1802) who apparently took the name from Commerson (pre Linnean), Jordan and Evermann (1896), present a history of Caranx which wi l l be quoted here, but they give the type of the genus incorrectly. The type is Scomber carangus Bloch = C, hippos (Linnaeus) by subsequent designation of Bleeker. (Lacepede gave no type), "The name Caranx was apparently recorded in manuscript by Commerson, who applied i t to Caranx speciosus (now Gnathanodon). In printed nomenclature Caranx was f i r s t used by Lacepede; who adopted the name from Commerson applying i t to a large group containing among other species trachurus. carangus, speciosus. ferdau, and ruber. This genus was further delimited by Rafinesque........next further restricted by Cuvier and Valenciennes" (Jordan and Evermann, 1896). The present restriction, and the one commonly used today, is that of Bleeker (1851), whose "type by subsequent designation is the correct type of the genus. A generic description of Caranx is included as in most of the previously published works the generic descriptions are based on too few specimens of each Caranx species (Jordan and Evermann, 1896; Meek and Hildebrand, 1925), or based on Caranx from different areas and do not conform to the Caranx of the tropical east Pacific (Williams, 1958). Some of the older Caranx descriptions do not follow Bleeker's restriction and include several other genera ( G i l l , 1863; Jordan and Gilbert, 1884; Gtinther, 1869; Weber and DeBeaufort, 1931). DESCRIPTION OP THE GENUS CARANX Caranx Lacepede (as restricted by Bleeker, 1851). Body ovate or oblong, fusiform to compressed, anterior-dorsal profile - 13 -sometimes strongly concave (as in large C. lugubris) to gently arched (C_. crysos and caballus). Adipose shield on eyes moderately to well developed. Pectoral-girdle not crossed by groove near isthmus. Mouth moderate to large, maxillary broad with well developed supplemental bone, extending to below eye. Premaxillary protractile. Teeth developed in one row on upper and lower jaws (two rows in C_, vine tus lower jaw), upper jaw with an inner band of fine teeth. Small teeth also present in patches on the vomer, palatine and tongue. Jaw teeth occasionally becoming deciduous or overgrown with flesh in the large specimens of some species. G i l l rakers moderate in size. Preopercle entire in adults, serrated in the very young, with a membraneous border. Dorsal spines low (l/2 height of elevated rayed dorsal) to moderate (slightly higher than elevated rayed dorsal), seven or eight in number connected with a membrane except the posterior two, and collapsing into a groove in the back. Often a small anteriorly pointed "spine" from the pterygiophore at the anterior end of the spinous dorsal (See Pig. 2), overgrown with skin in larger specimens. Rayed dorsal and anal fins falcate anteriorly except in C. vinetus. Two spines anteriorly separated from rayed anal. Never any dorsal or anal finlets. Ventral fins moderate, never extended, one spine and five rays. Pectoral fins long and falcate. Caudal f i n strongly forked, the peduncle slender, adipose ridges above and below lateral line half on peduncle and half on anterior caudal f i n . Scales small, cycloid, often irregular and sometimes deciduous. Breast scaled except in C. hippos where i t is naked except for a small patch anterior to the ventral fins. Lateral line arched anteriorly, the posterior part armed with strong lateral scutes, widest on the peduncle. Young often with vertical cross bars fading with age except in C_. vine tus where the bars are retained throughout l i f e . The placing of C. vine tus in the genus Caranx has been questioned by - 14 -Berry (pers com) using Williams' (1958) definition of Caranx. This species does not agree with Williams' definition because i t has two rows of teeth on the lower jaw and does not have falcate dorsal and anal fins. However, any attempt to place C. vinetus in another genus (probably Carangoides) presents more problems than are solved; the only way to keep to Williams' definition would be to erect a new genus for C» vinctus which seems undesirable. Therefore in this work Williams' definition of Caranx is broadened to include C. vinctus. - 15 -DESCRIPTIONS OF CARANX SPECIES CARANX CABALLUS GUNTHER (Fig. 6) Svnonvmv Trachurus boops Girard, 1858, p. 108 (desc. and fig.) - Not T. boops of C.&V. Caranx boops G i l l . 1862, p. 261 (desc.) - Not C. boops of C.&V. Caranx caballus Gunther. 1869, p. 431 (desc. and fig.) Jordan and Gilbert, 1884, p. 199 (syn.) Evermann and Jenkins, 1891, p. 138 (dist.) Eigenmann and Eigenmann, 1892, p. 353 (dist.) Jordan and Evermann, 1896a, p. 207 (dist.) Jordan and Evermann, 1896, p. 921 (desc.) Snodgrass and Heller, 1905, p. 333-427 (dist.) Meek and Hildebrand, 1925, p. 359 (desc. and fig.) Walford, 1937, p. 73 (desc. and f i g . and dist.) Hildebrand, 1946, p. 208 (desc. and fig.) Caranx girardi Steindachner. 1869, p. 25 Caranx crysos Nichols. 1920, p. 29 (part desc.) Caranx crysos caballus Nichols and Murphy, 1944, p. 243 (desc.) Material Examined Cat. No. Specimens W51-39 4 ¥51-49 1 W51-51 2 W51-56 1 W59-12 1 W59-248 1 BC54-46 3 BC54-52 1 BC55-38 2 BC56^337 1 BC57-94 1 BC57-95 1 BC57-98 4 BC57-104 1 BC57-108 1 BC57-144 2 BC57-168 1 Size Range Locality (mm.) Tenados I., Sinaloa, Mexico 212-220 Venados I., Sinaloa, Mexico 207 Venados I., Sinaloa, Mexico 285-294 Mazatlan, Sinaloa, Mexico 194 Point Willard, Lower California, Mexico 203 Pulmo, Lower California, Mexico 179 Acapulco, Guerrero, Mexico 186—191 Socorro I., Revillagigedo I *, Mexico 351 Zihautanejo, Guerrero, Mexico 175-181 Indefatigable I., Galapagos Is., Eq. 262 Petacalco Bay, Guerrero, Mexico 195 Petacalco Bay, Guerrero, Mexico 284 Petacalco Bay, Guerrero, Mexico 181-310 Santiago Bay, Colima, Mexico 343 Navidad Bay, Compilo, Mexico 200 Socorro I., Revillagigedo I ", Mexico 250-268 Maria Magdelena I., Tres Marias I s*, Mexico 271 - 16 -BC59-236 1 BC59-256 1 BC59-270 1 BC59-680 3 BC59-689 3 BC60-5 2 BC60-15 1 BC6CKL13 2 BG60-114 2 BC60-116 3 BC60-465 2 BC60-466 2 BC60-482 1 BC60-485 1 BC60-502 6 BC61-142 1 BC61-181 1 Cape San Lucas, Lower California, Mexico Clarion I., Revelligigedo I s*, Mexico San Juanito I., Las Tres.Marias I *, Mexico Chame Point, Panama Panama City Mkt., Panama Manjanillo Mkt., Colima, Mexico Acapulco Mkt., Guerrero, Mexico Tabago I., Panama Tabago I., Panama Tabago I., Panama Isla l a Granda, Guerrero, Mexico Isla l a Granda, Guerrero, Mexico Maria Magdelena I., Las Tres Marias Maria Magdelena I., Las Tres Marias Cape San Lucas, Lower California, Mexico I s* ,Mex, IS\Mex, Maria Magdelena I., Las Tres Marias I' Socorro I., Revellegigedo I s., Mexico Mex, 176 169 284 197-203 205-212 294-311 327 265-280 167-191 227-273 173-195 297-313 ,314 ,316 195-403 ,314 322 Description Dorsal VIII-I, 21-24 (one specimen 20 but with damaged fi n ) ; anal II-I, 18-21 rarely 17; caudal minor rays-dorsal 7-9, ventral 7-9; caudal major rays 17-21; pectorals I, 20-23; ventrals I, 5. Lateral scutes 38-51. Vertebrae 10 + 15 = 25. G i l l rakers on lower limb of f i r s t arch 25-31, usually 27-31, Body fusiform, not compressed, snout moderate not steep. Depth in length 3.32-4.78 (x 3.78). Head in length 3.49-4.01 (x 3.77). Pectoral f i n in length 2.60-3.57 (x 3.09). Upper jaw length in head 2,46-2.80 (x 2.68). Eye length in head 3.75-5.03 (x 4,32), this character varies greatly with size. Por 95$ ranges of proportions see Pig. 3, 4 and 5. Scales small, cycloid, somewhat irregular and deciduous, present on cheek, postorbital region and breast. Anterior arched portion of lateral line about half the length of the posterior straight portion. Spinous dorsal not quite as high as elevated portion of rayed dorsal f i n . Rayed dorsal and anal fins with obvious and loose scaly sheath. Ventral fins short, do not reach anus. Pectoral fins very long and falcate in adults, considerably shorter in young. Soft dorsal Caranx caballus B C 6 0 - 4 6 5 specimen No. 1 - 16b -Figur© 7 and anal fins falcate, their longest rays equal to between 1/4 and 1/3 of the length of the f i n base. Teeth present on both jaws, vomer, palatine and tongue, jaw teeth deciduous i n many larger specimens. Single row of teeth on lower jaw, single row with small teeth behind on upper jaw. Eye with adipose shield both anterior and posterior. Colour in Alcohol: Fins: Spinous and rayed dorsal dusky; caudal dusky; anal pale yellowish, some dusky area around edges; ventral pale yellowish; pectoral pale yellowish with a dark base on the underside. Operculum with black or very dark grey-brown spot (see Fig. 6). Body dark dorsally (irridescent blue in l i f e ) , ventrally pale yellowish (silver in l i f e ) . Distribution: (See Fig. 7) C, caballus is endemic to the tropical East Pacific area. The southward limit i s on the Peruvian Coast, probably Cape Aguja, (Walford, 1937). To the North i t has been taken at San Diego (Eigenmann and Eigenmann, 1892). It is known also from the offshore islands in the Revillagigedo group and on the Galapagos group (Snodgrass and Heller, 1905)* Specimens used in this study are taken from the areas shown on Fig. 7. CARANX VINCTUS JORDAN AND GILBERT (Fig. 8) Synonymy Caranx sp. Lay and Bennet, 1839, p. 55 (desc. and fig.) - 18 -Caranx vinetus Jordan and Gilbert, 1882, p. 349 (desc. and fig.) Jordan and Gilbert, 1884, p. 199 (syn.) Jordan and Evermann, 1896, p. 918 (desc.) Jordan and Evermann, 1896a, p. 207-584 (dist.) Gilbert and Starks, 1904, p. 77 (part desc.) Meek and Hildebrand, 1925, p. 358 (desc. and fig.) Walford, 1937, p. 74 (desc. and f i g . and dist.) Nichol, 1944, p. 124 (desc. and dist.) Carangus vinetus Jordan and Evermann, 1902, p. 305 (colour) No. Size Range Cat.No. Specimens Locality (mm.) W53-185 3 Acapulco, Guerrero, Mexico 92-103 ¥54-434 2 Locality unknown in Costa Rica 60-161 ¥58-46 4 S. of Topolobampo, Sinoloa, Mexico 231-266 W58-47 7 S. of Topolobampo, Sinoloa, Mexico 119-284 BC57-76 1 Acapulco Mkt., Guerrero, Mexico Port (Puerto) Arista, Oaxaco, Mexico 197 BC60-25 1 254 BC60-30 2 larras, Sinaloa, Mexico 168-172 BC60-31 2 2\ miles south of Mazatlan, Sinaloa, 197 Mexico Description Dorsal VIII-I, 22-25 (one specimen 21 but with damaged fi n ) ; anal II-I, 19-21; caudal minor rays-dorsal 8-11; ventral 8-10; caudal major rays 17-19; pectorals I, 18-20; ventrals I, 5. Lateral scutes 45-52. Vertebrae 10 + 14 = 24. G i l l rakers on lower limb of f i r s t arch 26-30 usually 28-30. Body fusiform, slightly compressed, snout moderate, not steep. Depth in length 2.84-3.34 (x 3.08). Head in length 3.49-3.84 (x 3.72). Pectoral f i n in length 2.99-5.00 (x 3.36). Upper jaw length in head.2.65-3.14 (x 2.77). Eye length in head 3.55-5.07 (x -4.12), this character varies greatly with size. For 95$ ranges of proportion see Figs. 3, 4 and 5. Scales small, cycloid, somewhat irregular, present on cheek, postorbital region and breast. - 19 -Anterior arched portion of lateral line about three quarters the length of the posterior straight portion. Spinous dorsal as high as elevated portion of rayed dorsal f i n . Rayed dorsal and anal fins with reduced scaly sheath on anterior portion of f i n only. Ventral fins moderate reaching to or slightly past anus. Pectoral fins very long and falcate in adults, considerably shorter in young. Soft dorsal and anal fins raised anteriorly but not falcate. Teeth present on both jaws, vomer, palatine and tongue, 2 rows of teeth on lower jaw, one irregular row with small teeth behind on upper jaw. Eye with adipose shield only moderately developed both anterior and posterior. Colour in Alcohol: Fins: Spinous dorsal dusky; rayed dorsal and anal pale yellowish; caudal yellowish; pelvic pale yellowish; pectoral pale yellowish, small dark area at base on the underside. Operculum with black or very dark grey-brown spot (See Fig, 8). Body dark dorsally (irridescent blue in l i f e ) , ventrally pale yellowish (silver in l i f e ) . Seven or eight dark vertical bars on the side of the body reaching to just below the midline. Distribution: (See Fig. 9) C. vinctus is endemic to the tropical east Pacific area. The published southward limit is Panama (exact location missing (Walford, 1937)). To the north i t is known from the southern part of the Gulf of California (Jordan and Evermann, 1896) and Conception Bay (Walford, 1937). The specimens used in this study are taken from the areas shown in Fig, 9. C a r a n x vinctus B C 6 0 - 3 0 specimen No.1 F i g u r e 8. F i g u r e 9, - 20 - -CARANX HIPPOS LINNAEUS (Pig. 10) Synonymy Scomber hippos Linnaeus, 1766, p. 492 (desc.) Carangus chrysos G i l l . 1863, p. 434 (desc.) Carangus hippos Grill. 1863, p. 434 Carangus hippus Jordan and Gilbert, 1883, p. 269 (misspelling) Caranx caninus Gunther. 1866, p. 602 (desc.) Gunther, 1869, p. 432 (desc.) Walford, 1937, p. 72 (desc. and f i g . and dist.) Caranx hippos Gunther. 1860, V 2, p. 449 (desc.) Jordan and Gilbert, 1883, p. 269 (dist.) Jordan and Gilbert, 1884, p. 200 (syn.) Jordan and Evermann, 1896, p. 920 (desc.) Gilbert and Starks, 1904, p. 77 (desc.) Starks, 1906, p. 762 (dist.) Starks, 1911, p. 48 (osteol.) Kendall and Radcliffe, 1910, p. 99 (dist.) Nichols, 1920a, p. 44 (desc.) Nichols, 1937a, p. 58 (desc.) Nichols, 1937, p. 1 (desc.) Hildebrand, 1939, p. 38 (dist. and canal) Nichols and. Roemhild, 1946, p. 15 (fin variation) Hildebrand, 1946, p. 208 (desc.) Smith, 1953, p. 218 (dist. S. Af.) Mather, 1954, p. 292 (dist. Atl.) Randall, 1955, p. 87 (dist. Pac. I s*) Robinson, 1959, p. 254 (dist.) Berry, 1959, p. 503 (dist. Atl.) Blanc and Bauchot, 1960, p. 488 (dist. W. Af.) Caranx carangus Gunther. 1860, p. 448 (desc.) Weber and DeBeaufort, 1931, p. 257 (dist. Indo Pac.) Caranx anthopygus Jordan. 1925, p. 16 Material Examined Cat. No. Specimens Locality Size Range (mm.) ¥51-22 BC56-142 BC56-145 BC57-78 BC57-87 BC57-118 16 1 1 2 6 2 Mazatlan, Sinaloa, Mexico Capoblanco, Peru Capoblanco, Peru Acapulco, Guerrero, Mexico Petacalco Bay, Guerrero, Mexico Chamela Bay, Colina, Mexico 146-289 550 444 196-198 108-172 345-380 - 21 -BC59-215 BC59-656 BC59-667 5 1 4 Mazatlan, Sinaloa, Mexico Cupica Bay to Orpu River, Columbia 30 M. N. of Panama City,-mouth Caimito River, Panama Santa Clara, Panama Los Angles, Panama Los Angeles, Panama Chiman Area, 90 M. S. Panama City Panama City Mkt», Panama Manzanilla, Coliraa, Mexico Puerto Arista, Oaxaco, Mexico San Bias, Nayarit, Mexico Acapulco, Guerrero, Mexico Maria Madre?, Tres Marias I s*, Mexico Maria Madre, Tres Marias I s*, Mexico Cape San Lucas, Lower California, Mexico 163-244 69 BC59-671 BC59-674 BC59-676 BC59-685 BC59-687 BC60-12 BC60-25 BC60-28 BC61-I26 N 906 3 2 1 4 2 2 2 1 1 1 1 1 113 93-111 355 109- 150 264-323 68-168 110- 114 N 907 N 708 143 123 508 468 730 Description Dorsal VII or VIII-I, 18-23 (usually VIII, 19-21); anal II-I, 15-18; caudal minor rays-dorsal 8-9, ventral 7-9; caudal major rays 17-19 the dorsal half of which are always 10; pectorals I, 18-21; ventrals I, 5. Lateral scutes 27-42. Vertebrae 10 + 14 = 24. G i l l rakes on lower limb of f i r s t arch 15-19 (usually 16-18). Body moderately compressed, snout steep but rarely i f ever concave in profile. Depth in length 2.47-3.74 (x = 2.92) varies with size of fish (see Pig. 3). Head in length 3.11-3.58 (x 3.36). Pectoral f i n in length 2.76-3.94 (x -3.20). Upper jaw length in head 2,16-2.55 (x -2.30). Eye length in head 3.28-5.77 (x 4.22), this character varies greatly with size. For 95$ ranges of proportions see Figs, 3, 4 and 5. Scales small, cycloid, somewhat irregular, present on cheek and postorbital region, those on cheek very small, absent from breast except for a small, diamond shaped, patch immediately anterior to the ventral fins. Soft dorsal and anal fins falcate, the longest rays of the fins equal to l/3 to l/2 the length of the f i n base. Anterior arched portion of lateral line about 4/5th the length of the - 22 posterior straight portion. Spinous dorsal about half as high as elevated portion of rayed dorsal f i n . Rayed dorsal and anal fins with reduced scaly sheath on anterior portion of the fins only. Ventral fins moderate reaching to the region of the anus. Pectoral fins very long and falcate in adults, considerably shorter in young. Teeth present on both jaws, vomer, palatine and tongue, becoming deciduous in large specimens. Single row of teeth some enlarged and canine like on the lower jaw, similarly on the upper jaw but with small teeth behind. Eye with adipose eye shield both anterior and posterior. Colour in Alcohol: Fins: Spinous dorsal dusky; rayed dorsal yellowish with dark edges and dark f i n rays on posterior 3/4 of f i n ; caudal yellowish with dusky edges; anal yellowish with dusky markings on edge. Ventral f i n yellowish; pectoral f i n pale yellowish with dark area at base on underside and dark spot on lower portion. Operculum with a black or very dark grey spot (See Fig, 10). Body dark dorsally (irridescent blue in l i f e ) , ventrally pale yellowish (silver in l i f e ) . Four or five vertical dark bars across body reaching to below the midline in young, disappearing in adults. Distribution (See Fig. l l ) C. hippos is regarded by several authors to be circumtropical (Berry, 1959; Briggs, I960). However, several fauna works claiming to include a l l Caranx species from the areas concerned omit (3. hippos. Most notable of these is Williams (1958) on tropical British East Africa, Randall's (1955) report on the fishes of the Gilbert Is. contains only a questionable reference to the presence of C_. hippos. The positive range may be placed as follows: both east and west coasts of tropical America, South African tropical waters Caranx hippos B C 5 7 - 8 7 specimen No.1 Figure 10. - 22b -F i g u r e 11. - 23 -(Smith, 1953), west coast of Africa (Blane and Bauchot, I960), Indo-Australian Archipelago (Weber and Debeaufort, 1931, Caranx carangus Bloch = C. hippos Linnaeus), and the Chinese Coast (Fowler, 1937a). It therefore may be considered world wide except for some small areas where C. hippos may not exist. In the tropical east Pacific i t is found from the extreme north of the Gulf of California (Robinson, 1959) and Cape San Lucas south to Cape Aguja, Peru. The specimens used in this study are taken from the areas shown on Fig. 11. CARANX MARGINATUS GILL (See Fig. 12) Synonymy Carangus marginatus G i l l . 1863, p. 166 (desc.) Jordan and Evermann, 1902, p„ 306 (desc.) Caranx latus = fallax = marginatus Jordan and Gilbert 1883 (dist.) Caranx latus = marginatas Jordan and Gilbert, 1884, p. 200 (syn.) Caranx latus Jordan and Eollman, 1890, p. 180 (dist.) Evermann and Jenkins, 1891, p. 138 (dist.) Jordan and Evermann, 1896a, p. 207-584 (dist.) Snodgrass and Heller, 1905, p. 333-427 (dist.) Caranx fallax (marginatus) Jordan and Gilbert, 1883a, p. 373-378 (dist.) Caranx marginatus Jordan and Evermann, 1896, p. 922 (desc.) Jordan and Evermann, 1896a, p. 207-584 (dist.) Gilbert and Starks, 1904, p. 78 (desc.) Fowler, 1905, p. 81 (desc. from Hawaii) Snodgrass and Heller, 1905, p. 333^427 (dist.) Kendell and Radcliffe, 1912, p. 99 (desc.) Meek and Hildebrand, 1925, p. 356 (desc. and fig.) Walford, 1937, p. 74 (desc. and fig.) Hildebrand, 1939, p. 38 Caranx sexfasciatus Nichols. 1938, p. 1 (dist.) Nichols and Murphy, 1944, p. 242 (desc.) Caranx hippos Giinther. 1869, p. 431 (in error) Material Examined No. Size Range Gat. No. Specimens Locality (mm.) ¥51-36 15 Esteso at Mazatlan, Sinaloa, Mexico 150-213 BC54-64 1 Cape San Lucas, Lower California, Mexico 128 BC56-444 1 Tower I., Galapagos I s* 140 BC56-445 1 Tower I., Galapagos I s* 87 BC56-449 3 Cocos I., Costa Rica, Mexico 91-101 BC56-454 ? 2 Cocos I., Costa Rica, Mexico 105-109 BC57-78 2 Acapulco, Guerrero, Mexico 241-243 BC57-175 1 Maria Magdalena I., Tres Marias Is;Mexico 186 BC58-383 3 Clarion I., Revillagigedo I s', Mexico 148-176 BC59-202 2 Maria Magdalena I., Tres Marias ISJ Mexico 106-116 BC59-241 5 Maria Magdalena I., Tres Marias I sJ Mexico 123-175 BC59-264 1 Cleopha I., Tres Marias I s*, Mexico 148 BC60-12 14 examined Manzanillo, Colima, Mexico 86-128 BC60-13 5 Acapulco, Guerrero, Mexico 100-22 BC60-17 5 Acapulco, Guerrero, Mexico 118-140 BC60-476 2 Chamela Bay, Nayarit, Mexico 94-97 BC61-125 1 Isla La Granda, Guerrero, Mexico 116 Description Dorsal VIII-I, 19-21; anal II-I, 15-17 rarely 14; caudal minor rays, dorsal 7-9, ventral 7-9; caudal major rays 17-19; pectorals I, 18-21, rarely 17; ventrals I, 5. Lateral scutes 28-36. Vertebrae 10 + 15 = 25. Grill rakers on lower limb of f i r s t arch 15-17. Body fusiform, moderately compressed, snout moderately steep never concave in profile. Depth in length 2.50-3.31 (x 2.89). Head in length 3.04-3.65 (x 3.40). Pectoral f i n in length 2,92-4.41 (x 3.43). Upper jaw length in head length 2.02-2.38 (x 2.19). Eye length in head length 3.23-4.29 (x 3.66), this character varies greatly with size. For 95$ ranges of proportions see Fig. 3, 4 and 5. Scales small, cycloid, somewhat irregular, present on cheek, postorbital region and breast. Anterior arched portion of lateral line slightly more than half the length - 25 of the posterior straight portion. Spinous dorsal about half as high as elevated portion of rayed dorsal. Rayed dorsal and anal fins with very reduced scaly sheath on anterior 1/3 of f i n only. Ventral fins reaching well beyond anus. Pectoral fins very long and falcate in adults; considerably shorter in young. Soft dorsal and anal fins falcate, the longest rays of the fins equal to 1/3 to 1/2 the length of the f i n base. Teeth present on both jaws, vomer, palatine and tongue, becoming grown over with flesh in some larger specimens, A single row of teeth on the upper and the lower jaws, a few anterior ones enlarged canine like, a patch of small teeth behind single row on upper jaw. Colour in Alcohol: Fins: Spinous dorsal dusky, rayed dorsal yellowish, dusky borders especially on the anterior portions of the f i n ; anal pale yellowish; caudal dusky yellow with darker posterior border. Ventral fins pale yellowish. Pectoral f i n yellowish, darker area at base on underside. Opercular spot pale or absent however there is a small black dot at the dorsal angle of the g i l l opening. Body dark dorsally (irridescent blue in l i f e ) , ventrally pale yellowish (silver in l i f e ) . Four to six dark vertical bars on sides of body extending to below the midline in young, disappearing with age. Distribution (See Fig, 15) C, marginatus is a part of the worldwide C. latus-sexfaseiatus»<narginatus complex (see page 42 ). Only the ranges of C. marginatus will be discussed here. C. marginatus is known from the tropical east Pacific and a questionable record from Hawaii (Fowler, 1905). It occurs from the south Gulf of California (Jordan and Gilbert, 1883a), to Panama (Kendall and Radcliffe, 1912) and on the offshore islands in the Revilligigedo group, Cocos I., Galapagos I s * Caranx marginatus ; BC59-241 specimen - No. 2 F i g u r e 12 - 25b Figure 13. - 26 -(Snodgrass and Heller, 1905). The areas where specimens used in this study were taken can be seen in Fig. 13, CARANX LUGUBRIS POEY (Fig. 14) Synonymy; Scomber ascensionis (non Osbeck) Bloch and Schneider, 1801, p. 33 Caranx ascensionis (non Osbeck) Cuvier in Cuvier and Valenciennes, 1833, p. 102 (desc. and fig.) Gunther, 1860, p. 432 (desc.) Caranx lugubris Poey, 1860, p. 22 (desc. and fig.) Jordan and Gilbert, 1882a, p. 227 (desc. and dist.) Jordan and Gilbert, 1884, p. 201 (syn.) Jordan and Evermann, 1896, p. 924 (desc.) Jordan and McGregor, 1899, p. 271-284 (dist.) Meek and Hildebrand, 1925, p. 352 (desc.) Walford, 1937, p. 76 (desc. and dist. and fig.) Woods in Shultz et a l , 1953, p. 514 (desc.) Randall, 1955, p. 88 (dist.) Berry, 1959, p. 523 (dist.) Blanc and Bauchot, 1960, p. 489 (desc.) Caranx tenebrosus Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka, 1927, p. 656 (desc. and dist.) Material Examined No. Size Range r + w Specimens Locality (mm.) BC54-53 1 Socorro I., Revillagigedo I s*, Mexico 288 BC54-403 2 Cocos I., Costa Rica 197-204 BC57-159 1 Socorro I., Revillagigedo I s', Mexico 310 BC58-383 1 Clarion I., Revillagigedo I s*, Mexico 229 BC61-179 13 Socorro I., Revillagigedo I s', Mexico 349-621 BC61-183 7 Socorro I., Revillagigedo I s*, Mexico 325-421 BC61-184 2 Socorro I., Revillagigedo I s', Mexico 354-372 BC61-187 1 Socorro I., Revillagigedo I s*, Mexico 755 - 27 -Description Dorsal VIII or VII-I, 20—22 (only occasionally VII spines); anal II-I, 17-19 (rarely 16); caudal minor rays, dorsal 8-9, ventral 8-9; caudal major rays 17-20; pectorals I, 19-22; ventrals I, 5. Lateral scutes 26-34. Vertebrae 10 + 14 = 24. G i l l rakers on lower limb of f i r s t arch 17-20. Body compressed, snout steep and often concave in profile. Depth in length 2.44-3.10 (x 2.88). Head in length 3.20-3.60 (x 3.39). Pectoral f i n in length 2.72-3.40 (x 3.02). Upper jaw length in head length 2.30-2.66 (x 2.49). Eye length in head length 3.44-6.10 (x 4.61), this character varies greatly within any one size group. For 95$ ranges of proportions see Figs. 3, 4 and 5. Scales small, cycloid, somewhat irregular, quite firm, present on cheek, postorbital region and breast. Anterior arched portion of lateral line about half the length of the posterior straight portion. Spinous dorsal about l/3 as high as the elevated portion of rayed dorsal f i n . Rayed dorsal and anal with very reduced scaly sheath on anterior portion of f i n only. Pectoral fins very long and falcate in adults. Rayed dorsal and anal fins very falcate, longest rays of the f i n equal to more than l/2 the length of the f i n base. Teeth present on both jaws, vomer, palatine and tongue, becoming overgrown with flesh or lost completely in large specimens. A single row of teeth on the upper and the lower jaw, these teeth subequal and often canine-like. Upper jaw with small teeth behind the main row. Eye with adipose shield both anterior and posterior. Colour in Alcohol: Fins: Spinous and rayed, dorsal dark brown almost black; anal, ventral and pectoral fins similar. Figure 14. - 27b -Figure 15 - 28 -No opercular spot. Body dark brown, almost black dorsally, ventrally not quite as dark but s t i l l dark brown. Lateral line scutes almost black. Distribution (Fig. 15) This species is noted to have an almost circumtropical distribution by Berry (1959) who records i t as occurring in the Western Atlantic (but not on the coast of the United States), Eastern Atlantic, Eastern and Western Pacific and Indian Ocean. Definite distribution records are from -the Indo-Pacific (Randall, 1955); from Hawaii and the East Pacific (Walford, 1937); from the West Coast of Africa (Blanc and Bauchot, I960), and from the Western Atlantic (Berry, 1959). However, the East African and Cape Area covered by Williams (1958) and Smith (1953) show no records. Weber and DeBeaufort (1931) have not recorded the fish from the Indo-Australian archipelago and Roxas and Agco (1943) do not mention C, luguhris from the Philippine Is. This fish appears to inhabit the rocky shores of islands rather than the mainland. For example, i t is present around islands in both the Pacific and the Atlantic, but is not recorded to date from the mainland of America. C. lugubris i s taken infrequently. The specimens used in this study were taken on the offshore islands of the Revilligigedo group and Cocos Is. (Fig. 15). CARANX MELAMPYGUS CUVIER (Fig. 16) Synonymyt Caranx melampygus Cuvier in Cuvier and Valenciennes, 1833, p. 116 (desc. and fig.) Giinther, 1860, p. 440 Jordan and Gilbert, 1882a, p. 230 (desc. and dist.) Jordan and Gilbert, 1884, p. 201 (syn.) Jordan, 1886, p. 361 (dist.) Jordan and Evermann, 1896, p. 925 (desp.) - 29 -Caranx stellatus Eydoux and Souleyet, 1840, p. 167 (desc.) Giinther, I960, p. 436 (desc.) Jordan, Evermann and Tanaka, 1928, p. 655 (desc. and dist.) Weber and DeBeaufort, 1931, p. 253 (desc.) Walford, 1937, p. 75 (desc. and f i g . and dist.) Roxas and Agco, 1941, p. 40 (desc. and dist.) Smith, 1953, p. 216 (desc. and dist.) Monro, 1955, p. 129 (desc. and dist.) Carangus melampygus Jordan and Evermann, 1902, p. 307 (colour) Snodgrass and Heller, 1905, p. 365 (desc.) Wakiya, 1924, p. 139-244 (desc. and dist.) Meek and Hildebrand, 1925, p. 353 (desc.) Weber and DeBeaufort, 1931, p. 248 (desc. and dist.) Fowler, 1936, p. 286 (desc. and dist.) Smith, 1953, p. 216 (desc. and dist.) Woods in Shultz et a l , 1953, p. 512 (desc. and dist.) Monro, 1955, p. 129~Tdesc. and dist.) Williams, 1958, p. 382 (desc. and dist.) Carangus forsteri Jordan and Evermann, 1905, p. 191 (not seen) Caranx forsteri Jordan and Seale, 1906, p. 230 Xurel melampygus Jordan. Evermann and Clark, 1930, p. 272 (dist.) Xurel stellatus Jordan, Evermann and Clark, 1930, p. 273 (dist.) Material Examined Cat. No. Specimens BC57-152 2 Socorro I.» BC58-383 18 Clarion I., BC59-261 4 Socorro I., BC59-264 4 Cleopha I., BC61-149 3 Maria Madra BC61-179 3 Socorro I., BC61-181 1 Socorro I., BC61-187 1 Socorro I., BC61-189 1 Socorro I., r S . Locality Revillagigedo Revillagigedo Revillagigedo Tres Marias I I., Tres Marias I Revillagigedo I * Revillagigedo I * Revillagigedo I * Revillagigedo I s * s. , Mexico , Mexico , Mexico Mexico s*, Mexico Mexico Mexico Mexico Mexico Size Range (mm.) 107-151 110-131 210-241 89-129 642-675 441-610 398 670 119 Description Dorsal VII or VIII-I, 21-23 (usually VIII-I, 22-23); anal II-I, 18-20 (one specimen at 16); caudal minor rays-dorsal 8-9, ventral 7-9; caudal - 30 -major rays 16-19; pectorals I, 18-21; ventral I, 5. Lateral scutes 29-37 (usually 33-36). Vertebrae 10 + 14 = 24. G i l l rakers on lower limb of f i r s t arch 18-19 (rarely 16-17). Body compressed, snout f a i r l y steep in young, to steep in large specimens, profile of head often concave in large specimens. Depth in length 2.47-3.19 (x 2.74) varying with size (See Fig, 4), Head in length 3.07-3.71 (x 3.52). Pectoral f i n in length 2.69-3.47 (x 3.13). Upper jaw length in head length 2.31-2.90 (x 2,53). Eye length in head length 3.32-7.84 (x 4,49), this character varies greatly with size. For 95$ ranges of proportions see Figs. 3, 4 and 5. Scales small, cycloid, somewhat irregular, well embedded, present on cheek, postorbital region, and breast. Anterior arched portion of lateral line about half the length of the posterior straight portion. Spinous dorsal from 1/2 to 3/4 as high as the elevated portion of rayed-dorsal f i n . Rayed dorsal and anal fins falcate, the longest rays equals from 2/5 to slightly more than l/2 of the length of the f i n base, scaly sheath reduced and only present on anterior half of fins. Ventral fins moderate in length reaching past the anus. Pectoral fins long and falcate in a l l specimens studied. Teeth present on both jaws, vomer, palatine and tongue, jaw teeth becoming overgrown with flesh or deciduous in some large specimens. A single row of teeth on the lower and the upper jaws; the upper with patch of small teeth behind. Colour in Alcohol: Specimens under 150 mm,: Spinous and rayed dorsal dusky with darkened borders on rayed dorsal; anal dusky except anterior falcate lobe which is dark. Ventral fins light with faint dusky tint at distal end; pectoral fins light; caudal dusky with darker borders. - 31 -Opercular spot absent. Body dark dorsally (irridescent blue in l i f e ) , ventrally silver as in l i f e , no snots present. Specimens 200-300 mm.: Spinous and rayed dorsal dark, anterior portion of rayed dorsal almost black; anal dark, anterior lobe being almost black. Ventral fins dark on lateral edge and nearing distal end, light medially and proxmally; pectoral fins light, slightly darker than young; caudal darker than young but not nearly black. Opercular spot absent. Body dark dorsally (irridescent blue-green in l i f e ) , ventrally dirty silver as i n l i f e ; a few spots (10—20) present on each side of the body (Fig. 15), Specimens greater than 350 mm,: Fins as in mid-sized specimens except a l l fins a l i t t l e darker and the caudal becoming very dark brown with lighter posterior margin. Opercular spot absent. Body very dark (almost black) dorsally (irridescent blue-green on dark backgroundin l i f e ) , ventrally a golden shade as in l i f e , many spots on side of body and head. From the above i t can be seen that the colour pattern of C, melampygus changes radically with growth, which has led many authors to believe the young and the adults to be distinct species (See Table III). Distribution: (Fig. 17) Caranx melampygus is reported from the tropical Indian Ocean and the Red Sea (Smith, 1953; Williams, 1958), throughout the Indo-Pacific region (Weber and DeBeaufort, 1931; Randall, 1955; Monro, 1955) and in the east Pacific. This species is not recorded from the Atlantic Ocean, In the east Pacific i t is recorded from the Tres Marias I s* south to Panama and on the offshore islands of the Revillagigedo group and Cocos (Walford, 1937). Figure 16. - 31b -Figure 17. - 32 -CARANX MEDUSICOLA JORDAN AND STARKS Synonymy: Caranx medusicola Jordan and Starks, 1895, p. 430 (desc) Jordan and Evermann, 1896, p. 924 (desc. and f i g . and dist.) Gilbert and Starks, 1904, p. 78 (desc. and dist.) Meek and Hildebrand, 1931, p. 355 (desc.) Walford, 1937, p. 75 (desc. and dist.) Carangus medusicola Jordan and Evermann, 1902, p. 306 Material Examined No. Cat. No. Specimens Locality Size Range (mm.) BC54-64 2 Cape San Lucas, Baja California, Mexico 112-114 BC60-488 1 Maria Magdalena I L a s Tres Marias I s*, Mexico 157 Description Dorsal VIII-I, 21-23; anal II-I, 19-20; caudal minor rays-dorsal 7-8, ventral 7-8; caudal major rays 18; pectorals I, 20; ventrals I, 5. Lateral scutes 37-40. Vertebrae 10 + 14 = 24. G i l l rakers on lower limb of f i r s t arch 19. Body compressed, snout f a i r l y steep, profile straight, not concave on the three specimens examined. Depth in length 2.45-2.59 (x 2.52). Head in length 3.58-3.66 (x 3.62). Pectoral f i n in length 3.70-5.20 (x no meaning). Upper jaw length in head length 2.43-2,64 (x 2,54), Eye length in head length 4,2-4.3 (x 4.25). Scales small, cycloid, somewhat irregular not embedded, present on cheek, postorbital region and breast. Anterior- arched portion of lateral line about half the length of the posterior straight portion. Spinous dorsal from 3/4 to equal to the height of the elevated portion of the rayed dorsal. Rayed-dorsal and anal fins - 33 -falcate, but anal rounded on falcate end. The longest dorsal ray equal to 1/3 to 1/2 of the length of the base, the longest anal ray equal to less than 2/5 the base of the f i n . Scaly sheath reduced, present only on anterior half of f i n bases. Ventral fins moderate in length reaching past the anus. Pectoral fins long and falcate in specimens studied. Teeth present on both jaws, vomer, palatine and tongue. A single row of teeth on the lower and the upper jaws. The upper with patch of small teeth behind. Colour in Alcohol: Pins: Spinous and rayed dorsal quite light with darker borders except the anterior-falcate lobe which is dark, almost black, Ventrals light; pectoral light; caudal dusky with darker borders. Opercular spot absent; body dark brown dorsally, ventrally yellowish, no spots present. Distribution: (Pig. 19) The known distribution of this species is from Mazatlan (Jordan and Evermann, 1896) to Panama (Gilbert and Starks, 1904), and i t is reported by these authors to occur commonly. However, many specimens supposedly of this species may be C. melampygus. The area in which i t is reported to be commonly found is well represented in our collections of Caranx and yet we have only three specimens of C. medusicola. Mr, F. Berry, on checking some of the collections at U.C.L.A., feels that many of the identified C, medusicola are in fact C, melampygus. A discussion of the differences in these species is on page 45 « The specimens I have examined come from Cape San Lucas and the Tres Marias Islands 0 5 cw. C a r a n x ) m e d u s i c o l a B C 6 0 - 4 8 8 s p e c i m e n N o . 2 Figure 18. - 33b -Figure 19. - 34 -CARANX sp. BC56—434 1 specimen Indefatigable I., Galapagos I * This specimen from the Galapagos Islands does not f i t any available key to the species. The specimen is superficially similar to C_. marginatus but the vertebrae are 10 + 14 = 24 whereas C. marginatus has 10 +15 =25. Also the upper jaw length in head ratio is much higher in Caranx sp. than in C. marginatus (2.52 in Caranx sp. and 2,02-2.38 (x 2.19) in C, marginatus). Some of the characters of this form are as follows. Fork length, 68 mm.; depth, 24.7 mm,; head length, 19..9 mm.; pectoral f i n length, 14,9 mm,; upper jaw length, 7.9 mm.; eye diameter 4.9 mm. Depth in length 2.75; head in length 3.42; pectoral f i n in length 4.56; upper jaw in head, 2,52, Dorsal VIII-I, 20; anal ?II-I, 16; pectoral I, 19; ventrals I, 5. G i l l rakers on lower limb of f i r s t arch 18. Vertebrae 10 + 14 = 24. Caudal scutes 30. Six distinct dark vertical bands on side of body, extending almost to the ventral surface. Opercular spot present. - 35 -COMPARATIVE MATERIAL FROM ATLANTIC As well as the material previously mentioned, specimens of C» latus and C. crysos from the Atlantic were examined. CARANX LATUS AGASSIZ Material Examined No. Size Range Cat. No. Specimens Locality (mm,) BC60-2 1 Sacraficius I., 7 Vera Cruz, Mexico 67 UMML138 1 Plantation Key, Florida 112 UMML1575 2 Bimini, Bahamas 53 UMML1749 1 Bahia de legua, Puerto Rico 160 Description Dorsal VIII-I, 20-21; anal II-I, 16-18; pectoral I, 18-20; ventrals I, 5. Lateral scutes 33-38. Vertebrae 10 + 14 = 24. G i l l rakers on lower limb of f i r s t arch 16-17. Depth i n length 2.65-2.84. Head in length 2,67-3.31. Pectoral f i n in length 3,33-4,22. Upper jaw in length in head 2.03-2,65. Eye length in head 3,19-3,84, A discussion of the relationship of C, latus and C. marginatus can be found on page 42 . CARANX CRYSOS MITCHELL Material Examined - 36 -No. Cat. No. Specimens Locality Size Range (mm.) BC58-398 BC61-92 UMML408 UMML1150 UMML1179 UMML1457 UMML2150 UMML3048 UMML7397 UMML7784 1 1 1 1 1 1 1 1 1 1 Little Duck Key, Florida East Coast of Florida Florida Bay, Monroe, Florida Key Largo, Florida 300, 16• N; 80°, 21« W-Combat Stn. # 70 Virginia Key, Florida Virginia Key, Florida Dry Tortugas, Florida Dry Tortugas, Florida Long Key, Florida 166 100 178 100 146 149 184 175 156 91 Description Dorsal VIII-I, 22-24; anal II-I, 19-f20; pectoral I, 20-24; ventrals I, 5. Lateral scutes 45-51. Vertebrae 10 + 15 = 25. Grill rakers on lower limb of f i r s t arch 26-28. Depth in length 2.53-3.49 (x 3.20). Head in length 3.51-3.75 (x 3.62). Pectoral f i n in length 3.05-5.25 (x 3.69). Upper jaw length in head 2,24-2.53 (x 2.42). Eye length in head 3.48-4.46 (x 3.91). A discussion of the relationships of C. crysos and C. caballus can be found on page 39 • KEY TO THE SPECIES OF CARANX IN THE TROPICAL EAST PACIFIC There are seven known species of the genus Caranx in the tropical east Pacific. The keys in Meek and Hildebrand (1925), Jordan and Evermann (1896), and Walford (1937), do not adequately differenciate these species. Scute counts, anal and dorsal f i n ray counts and colour pattern are extensively used in these keys. Overlap was found in a l l such characters for Caranx marginatus, melampygus and medusicola. The key given below separates a l l the species of Caranx known from the tropical east Pacific. A Grill rakers on lower limb of f i r s t arch 22 or more (including rudiments). B Body with dark bars; scaly sheath on dorsal and anal fins reduced, restricted to anterior portion only. Caranx vinetus BB Body without dark bars; scaly sheath on dorsal and anal fins well developed - continuous along f i n Caranx caballus AA Grill rakers on lower limb of f i r s t arch less than 22 (including rudiments)• C Scales absent from breast except small diamond shaped patch in front of ventral fins. CC Breast scaled. D Body a l l dark brown; front of head usually concave; anterior portion of dorsal and anal f i n reaching well past l/2 f i n base. DD Body not a l l dark brown; front of head usually not concave; anterior portion of dorsal and anal f i n not reaching well past 1/2 f i n base. E Head length over upper jaw length 2,00-2.40 (usually between 2.05-2.25); body never spotted; g i l l rakers 17 or less, rarely 18; dorsal times anal f i n rays less than 365; vertebrae 25. Caranx hippos Caranx lugubris Caranx marginatus - 38 -EE Head length over upper jaw length 2.30-2.80 (usually 2.45-2.65). Body may be spotted in large individuals} g i l l rakers, 18 or more, rarely 17 or less; dorsal times anal f i n rays greater than 365; vertebrae 24. Anal fins pointed or almost so, longest anal ray 2\ or less in base of f i n ; anterio-dorsal profile not straight. Caranx melampygus FP Anal f i n round on anterior raised portion, longest anal ray greater than 2i in the f i n base; anterio-dorsal profile almost straight. Caranx medusicola DISCUSSION DEFINITION OF THE TROPICAL EAST PACIFIC The tropical east Pacific area is the region of tropical waters located between Magdalena Bay (approximately 24°N) and Cape Aguja (approximately 6°S) except during periods of the "El Nino" when the tropical waters extend further south to 12°S (Sverdrup, Johnson and Flemming, 1942), Included in this region are the offshore islands of the Revillagigedo group, Cocos and Clipperton Islands, the Galapagos Islands and perhaps Gaudalupe Island. The northern boundary of the tropical east Pacific region is a disputed question. It is placed as far north as the United States-Mexico boundary by Eckman (1953) and as far south as 23°N by Newell (1948). However, i t is generally accepted, on both a faunal and a temperature basis (20°C winter isotherm), that the area of Magdelena Bay can be considered as the boundary. Within the Gulf of California there appears to be a change from a completely tropical fauna in the south to a mixed tropical-warm temperate fauna in the north (Briggs, 1955). Tropical forms of Caranx are not known to exist south of Cape Ajuga, the southern limit of tropical waters. C. caballus is reported from as far north as San Diego (approximately 32° 35*N) and C, hippos is reported from the extreme north of the Gulf of California. Caranx then do move into the warm temperate waters to the north of the tropical zone but only infrequently. SPECIES AND SPECIES COMPLEXES Caranx caballus : crysos : fusus This complex of forms exists in the tropical waters of the east Pacific (caballus), west Atlantic (crysos) and subtropical Mediterranean (fusus). Tortonese (1952) places C. fusus and C. crysos in synonyme both as C. crysos. - 40 -The present author has examined C,, crysos and C, caballus and found both to have 25 vertebrae. Berry (pers. com.) has examined C. fusus and C_« crysos and has found these forms to have 25 vertebrae. The differences in some of the characters of C. crysos and C. caballus are shown i n the Table below. TABLE II L D L H H Lateral G i l l M Scutes Rakers C. crysos 3.20 3.62 2.42 46.7 26.8 2.53-3.49 3.51-3.75 2.24-2.53 45-51 26-28 C. caballus 3.78 3.77 2.68 44.1 28.6 3.32-4.78 3.49-4.01 2.46-2,80 38-51 25-31 Average range Average range L L H Using a character index on the above proportions (p + ^  + jjj-), 100$ separation is obtained, C. crysos having an index range from 8,6-9,5 and C. caballus having a range from 9.6-11.5. A graphical presentation of the proportional characters and index can be seen in Figs, 20 and 21. C. crysos and C. caballus have been synonymized as C. crysos by Nichol (1920) and as C. crysos crysos and C, crysos caballus by Nichol and Murphy (1944). Because 100$ separation between the two forms can be made by the use of a simple index a synonymy or subspecific synonymy is unwise and the two species should be retained as such. C. caballus i s endemic to the tropical east Pacific and C. crysos to the Atlantic; there are no closely related forms in the Indo-Pacific. As the isthmus of Panama was submerged in pre-pleistocene times (1-3 million years ago) i t is very likely that the C. crysos-caballus stock was - 40a -I 5 n crysos I N D E X caballus Figure 20. Character index of |f + - + S separating n D M C. crysos and C caballus. (C. crysos 91 mm to 184 mm C_j_ caballus 167 mm to 403 mm). h e a d l e n g t h f o r k l e n g t h 3 0 - i u p p e r j a w l e n g t h h e a d l e n g t h f o r k l e n g t h d e p t h 2 0 -£ 1 0 -o> E o <u o_ cn o- ZL i—r C c a b a l l u s C c r v s o s JEL 3 5 2 0 " 1 0 -0 -C . m a r g i n a t u s C . l a t u s 2 . 0 2 . 5 3 . 0 "ST 4 . 0 2 . 5 Figure 21. 3 . 0 i 3 5 4 . 0 4 . 5 - 41 -continuously distributed in both the east Pacific and Atlantic during this time. The evolution of these forms probably took place in the areas in which they exist today. The differentiation of C, caballus from C. crysos must have occurred since the last emergence of the Panama isthmus. Caranx vinctus This species is endemic to the tropical east Pacific and appears to have no very close relative in this or any other tropical region, Jordan and Gilbert (1882) in the original description of this form claim i t to be closely related to Caranx cibi Poey, C. cibi has now been synonomized with C_. bartholomaei Cuvier (Duarte-Bello, 1959). C, bartholomaei is a species which is found in the west Atlantic and Caribbean. Prom the descriptions of C_. bartholomaei in the literature and from the single specimen examined there would appear to be no close relationship between C, vinctus and C. bartholomaei. There exist large differences in dentition, g i l l raker count and scute count. C. vinctus does not f i t Williams* (1958) Caranx generic restriction as i t has two rows of teeth on the lower jaw and does not have falcate dorsal and anal fins. The origin of C, vinctus is d i f f i c u l t to ascertain. This species could be an example of a form originating in the Indo-Pacific and being displaced to the east Pacific since the last rising of the isthmus. An equally tenable argument is that C. vinctus evolved in the east Pacific-Atlantic region and was not able to survive the postulated low temperature in the west Atlantic during the pleistocene glaciation (see page 48 ) . Caranx hippos This species is known from almost a l l tropical waters. It has on occasion been named as a separate species in the tropical east Pacific, Caranx caninus (Gunther, 1869), The only recent author to follow Gunther - 42 -is Walford (1937). Gilbert and Starks (1904) have listed the slight differences between C. hippos on both sides of the Panama isthmus. East Pacific forms have a slightly higher g i l l raker count, more scutes on the lateral line and shorter ventral fins, but there is a great area of overlap in a l l cases. Nichols (1937) separates C, hippos on either side of the isthmus as subspecies, but the characters he used do not appear to be valid. He claims scute count on the Pacific form to be 33-40 and on the Atlantic form to be 27-33. The specimens examined for this study from the Pacific varied from 27-41. Another character Nichols used in his study was head length over eye diameter, a character which varies greatly with the size of the specimens (see page 4), Hildebrand (1939) states that C_. hippos is found in the Panama Canal locks but does not pass through the Canal. The young of C_. hippos is known to enter fresh water and the possibility exists that C. hippos does, on occasion, pass through the Canal. Meek and Hildebrand (1925) compared 50 specimens from the Atlantic coast and 70 from the Pacific coast and found no differences of sufficient magnitude to separate these populations as species. The conclusion reached from the above evidence is that C, hippos is the same species on both sides of the Panama isthmus but there are some slight morphological differences in the species between these areas. Caranx marginatus : latus s sexfasciatus The f i r s t problem encountered in this complex is the identification of C_. sexfasciatus. To give an example, Nichol and Murphy (1944) equate C. sexfasciatus with C, marginatus; Randall (1955) equates C, sexfasciatus from the Gilbert Islands, with C, hippos from Florida, and Woods (in Schultz et a l , 1953) describes C, sexfasciatus from the Marshall and Mariannes Island as a different form making i t equal to £. sexfasciatus of Quoy and Gaimard, - 43 -Prom the examination of two specimens of £. sexfasciatus Quoy and Gaimard, taken at Taihae (BC62-255) a vertebral count of 24 was obtained, one lower than C_. marginatus. As C. sexfasciatus. i f i t is a valid species, has not been collected from the tropical east Pacific the present work will deal only with C. marginatus and C. latus. C. marginatus is an east Pacific form and C, latus is a west Atlantic form. The proportional measurements and counts made on these two species with the exception of vertebrae are very similar. Figure 21 and the Table below summarize some of the data. TABLE III Dorsal Anal Lateral G i l l Vertebrae Rays Rays Pectoral Scutes Rakers c. latus 10 + 14 20-21 16-18 I, 18-20 33-38 16-17 c. marginatus 10 + 15 19-21 15-17 I, 18-21 28-36 15-17 The characteristics used to separate C. latus and C. marginatus by Meek and Hildebrand, namely eye diameter and body depth, do not appear to be valid. Eye diameter varies greatly with size (see page 4) and body depth into length has complete overlap (see Fig. 21). Vertebrae number separates these two species. As a l l marginatus examined had 25 vertebrae and a l l latus 24 vertebrae and in a l l other Caranx species examined the vertebral count was constant within a given species, i t is concluded that C. latus and CJ. marginatus are valid species. £, marginatus and C. latus have become differentiated since the last emergence of the Panama isthmus. Previous to the emergence they must have been continously distributed on both sides of the Americas. - 44 -Caranx lugubris This species i s distributed circumtropically around rocky islands. Often, as in the Americas, i t is not found on the mainland but only around islands. It appears to be quite constant in character throughout its range. As (3. lugubris is commonest in the Indo-Pacific region and as there are more islands here than in other tropical regions i t is suspected that this region is the probable area of origin. Caranx melampygus = stellatus Caranx melampygus and C, stellatus have been described as separate species by several authors, the division based on colour pattern, especially the lack of spots (C. melampygus) or presence of spots (C. stellatus). TABLE IV Number of Spots and Size of C. melampygus. Spots on 1 Side of No. Body Head Fish Size Specimens Area 0 0 89-129 mm. 4 0 0 107-151 mm. 2 0 0 110-131 mm. 18 0 0 119 mm. 1 10-30 1 or 2 210-241 mm. 4 50-100 100 396-670 mm. 5 50-100 100 642-675 mm. 3 r S . Cleopha I., Tres Marias I" Socorro I., Revillagigedo I" Clarion I., Revillagigedo I s* Socorro I., Revillagigedo I Socorro I., Revillagigedo I s] Socorro I., Revillagigedo I Maria Madre I.,Tres Marias I From the examination of the specimens above (from 89-675 mm.) i t is seen that the presence or absence of spots depends upon size. It is concluded that C. melampygus and C. stellatus are synonymous and must be called _C. melampygus. C. melampygus is a pan-Pacific form which i s absent from the Atlantic, On this basis i t is likely the C. melampygus has reached the east Pacific since the emergence of the Panama isthmus. If this is so, C, melampygus illustrates Briggs (1961) hypothesis that there i s at present a slow invasion of the tropical east Pacific by Indo-Pacific species of shore fish, which are crossing the east Pacific sea barrier. Caranx medusicola and melampygus Caranx medusicola has often been confused with C. melampygus. a species which i t very closely resembles. Differences between these species include; C. medusicola has a rounded falcate anal while C. melampygus has a pointed falcate anal, the length of the longest anal ray in C, medusicola divides into the anal f i n base more than 2.5 times while in C. melampygus the longest anal ray divides in the base 2.5 times or less. The anterior profile of the two species are quite distinct, that of C. medusicola being almost straight while that of C. melampygus is concave. , m i - — « i C m e l a m p y g u s 2 1 0 m m . / ' C , m e d u s i c o l a 1 5 7 m m F i g u r e 2 2. \ - 46 -It is very probable that C, medusicola is a valid species which is rarely collected (there are only 3 specimens in the U.B.C. collections) and i t is considered as such in this work. The origin of C. medusicola is di f f i c u l t to ascertain. Morphologically i t is very similar to C_. melampygus and on this basis a close phylogenetic relationship can be postulated. On zoogeographical evidence however, no close relationship is indicated. The reason for this i s : C. melampygus is pan-Pacific and because of its absence from the west Atlantic, appears to have reached the east Pacific since the last emergence of the Panama isthmus. C. medusicola is endemic to the east Pacific region. If C, medusicola and _C. melampygus are close phylogenetic relations, as is indicated by their morphological similarity, one of three possibilities exist. C. medusicola was once an Indo-Pacific form which was displaced to the eastern Pacific periphery and does not now exist in the Indo-Pacific; C. medusicola has evolved in the east Pacific from C_« melampygus very recently; or C. melampygus and C« medusicola in the past existed in the Pacific and Atlantic and todays distribution is the result of both these species failing to survive in the Atlantic. BARRIERS TO DISTRIBUTION Panama Isthmus Any comparisons of the Atlanto-Pacific fish faunas must take into account the existence and submergence of the Panama Isthmus. Stokes (i960) estimates submergence took place in the early Cenozoic, probably Paleocene or early Eocene and lasted until late Pliocene. Lindberge (i960) in his investigation of levels of the sea postulates that during the Sangamon interglacial period, previous to the Wisconsin glaciation, the marine water levels were 80 meters higher than at present, Flint (1957) states the - 47 -highest interglacial elevation in sea levels was over 30 meters. He shows further that around Santa Barbara the level 90 f t . above the present sea level is associated with marine fossils which are probably late Pleistocene, somewhat more than 30,000 years old. There are three areas today where the coastal lowlands extend from coast to coast, namely, in Nicaragua, in Panama and in south Mexico. It is therefore a possibility that during one of the interglacial periods, probably Sangaman, there existed a channel separating North and South America which would allow exchange in faunal elements between the Pacific and Atlantic Oceans. Today transport across the Isthmus is impossible for strictly marine forms as the Panama Canal runs through two bodies of fresh water, one of these 30 miles long (Lake Gatun), the other 3-4 miles long. Fish that can tolerate fresh or brackish water for moderate lengths of time could cross, however. East Pacific Barrier This barrier is not land but rather sea, a very large stretch of open water with no islands where shore fish or shore-bound pelagic fish could survive. The size and extent of this barrier may be seen in Fig. 23. The efficiency of the barrier has been estimated as 86.3$ at the specific level; i.e. 13.7$ of the total number of known shore fish have been able to cross in a west to east direction (Briggs, 1961), An elevation in the water level such as that proposed by Lindberge (i960) or Flint (1957) would submerge some of the low atoIs in the south Pacific and have the effect of increasing the size of the east Pacific barrier. A lowering of the sea levels as postulated during the Wisconsin glacial period would have no effect, as a drop in sea levels of 200 meters would only increase the size of the present islands in the tropical east Pacific and add no new islands in the areas of the east Pacific barrier (Data from U.S.Navy Hydrographic Office, 1961). Q > c o r a l r e e f a r e a s Figure 23. EQUATORIAL PACIFIC O C E A N Barrier area indicated by l a b e l . - 48 -Temperature Barriers At the present time the distribution of Caranx and other marine shore fishes in the tropical east Pacific is limited to the north and south by-temperature barriers. Distribution to the east and west is limited not by temperature but rather the previously discussed land and sea barriers. In Pleistocene and perhaps Pliocene times the eastern, northern and southern barriers may have differed from the present locations. Ewing and Donn (1956 and 1958) have postulated that the temperature in the Gulf of Mexico and Caribbean was considerably lower during the time of maximum ice melt in the interglacial periods. If this is the case, the existence of a channel between North and South America during the Sangaman interglacial may have had l i t t l e effect on Caranx distribution due to the presence of a low temperature barrier. Hubbs (1948 and I960) presents the hypothesis that the tropical east Pacific was cooler; marine temperatures decreasing by 3 C° in the winter and 8 C° in the summer. Such a drop in temperature would narrow the band of tropical waters. Hubbs also postulates that the paucity of the tropical east Pacific shore fishes is due to the aforementioned low temperature removing many species from this area, Briggs (1961) disagrees with Hubbs and postulates no catastrophic removal of shorefish species due to temperature. Briggs shows in Figures 1 and 2 of his recent (1961) paper that the 3 C° winter drop would only decrease the band of tropical waters by about 1/3, but he does not show the 8 C° summer drop postulated by Hubbs. The 8 C° reduction in temperature would reduce the tropical zone by l/2 its present width and cause some northward shift (See Figs. 24, 25 and 26). According to Briggs, there are two reasons for the paucity of the east Pacific tropical fauna; the f i r s t - 48a -Figure 25. Distribution of tropical waters with a winter temperature drop of 3 C & (From Briggs, 1961). - 48b -- 49 -is the effectiveness of the east Pacific barrier and the Panama isthmus barrier and the second is the lack of habitat in the east Pacific for coral reef fishes. Evidence for or against a marine temperature drop in the tropical east Pacific during the Pleistocene is sparce. Some of the evidence is reviewed here. Dillon (1956) following botanical and snow line evidence estimates land temperatures were 5 F° (2.8 C°) lower at the equator and 10 F° (5.6 C°) lower at 35°N than present temperatures. Interpolating from these estimates a drop in land temperature in the proximity of 7.5°P (4,1°C) could be expected in the region of the tropical east Pacific. Hubbs (1948) shows a close correlation between land and sea temperatures; therefore the drop of 4.1°C land temperature would be reflected in marine temperatures, but not quite to the same extent. Dorf (i960) presents more severe temperature decline. Working in paleobotany, Dorf estimates land temperatures in Central America during Pleistocene to be sub-tropical and warm temperate, there being no tropical zone. Arrhenius (1952) finds that along the line of 130°W the temperature was approximately 3 to 4 C° lower during glaciation. This work is based on Oxygen^g examination of core samples using planktonic foraminifera. Most evidence points to a decline in the marine temperatures during the periods of maximum glaciation. A drop of from 3 to 5° C appears to be the likely order of magnitude. - 50 -CARANX; ZOOGEOGRAPHY AND EVOLUTION The barriers, the tropical zone and the distribution of Caranx have been discussed previously. This information will now be speculatively examined in relation to evolution and variation in Caranx. and an evaluation of Hubbs' (i960) "temperate decrease theory" and Briggs' (1961) "barrier theory". There are three possible patterns of evolution and distribution to those Caranx forms endemic to the tropical east Pacific, Their patterns are: f i r s t l y , the endemic species have evolved within the tropical east Pacific since the last emergence of the Panama Isthmus; secondly, they have arrived in the tropical east Pacific since the last emergence of the Isthmus, coming from the Indo-Pacific where they do not exist today; or lastly they existed in the tropical east Pacific while the Isthmus was open but did not enter the Atlantic, or i f they did enter they did not survive there. Table V indicates the distribution of those species which are not endemic to the east Pacific and those which have close relatives across the Isthmus, As is seen from this Table, some forms vary to the extent of becoming different species in different areas of their ranges, while others are very similar throughout a l l the areas of their ranges, Caranx caballus and Caranx marginatus have most likely evolved in the tropical east Pacific since the emergence of the isthmus as both have a twin species on the Atlantic side of the isthmus. The theories on the paucity of the tropical east Pacific fauna previously mentioned (Briggs and Hubbs), will now be evaluated in terms of Caranx distribution. As circumtropical Caranx species (C. hippos and C. lugubris), add l i t t l e to either side of the argument, they will not be used. Five TABLE V Distribution and differentiation of non-endemic Caranx species in the east Pacific. INDO-PACIFIC EAST PACIFIC BARRIER EAST PACIFIC ISTHMUS WEST ATLANTIC DIFFERENTIATION C. sexfasciatus C. marginatus C. latus i C , marginatus from C . and C. sexfasciatus -on vertebral count latus 100$ C. caballus C. crysos 100$ with index C. hippos C. hippos C. hippos Slight variation C. melampygus C. melampygus Similar C. lugubris C. lugubris C. lugubris Similar - 52 -other Caranx species will be discussed, namely, C. caballus. C_. marginatus, 9.* vine tus. C_. melampygus and C, medusicola. Only one of the five east Pacific species (C. melampygus) is found in the Indo-Pacific. Conversly only one of the 12-20 (depending upon author) Indo-Pacific species (C. melampygus) is found in the east Pacific. We may say therefore, that the east Pacific barrier is 80$ effective for east to west movement and from 92$-95$ effective for west to east movement, in regard to the genus Caranx, The situation across the isthmus differs. Two of the five species in both Pacific and Atlantic are closely related (C, caballus : crysos and C. marginatus : latus). The Panama isthmus i s , therefore, only 60$ effective when considering either east-west or west-east movement in the past. Today, however, i t is 100$ effective as there are no identical species except the circumtropical ones. If Hubbs' theory is correct, then we may assume that the Caranx on either side of the isthmus were displaced during the Pleistocene and only in recent times have come back to either side of the isthmus. Also, the east Pacific species present today are largely due to re-invasion from the Indo-Pacific, Re-invasion from the Atlantic is impossible as the isthmus has blocked passage since the Pleistocene. Caranx does not show such a pattern. The affinities with the Indo-Pacific are very low, the affinities with the Atlantic somewhat higher. The present evidence indicates that the Pleistocene marine temperature was 3 to 4C° lower than i t is today, Hubbs postulates a 3°C lower winter temperature and an 8°C lower summer temperature. It is unlikely that the later (8 C°) lowering is correct as this would make the tropical zone - 53 -narrower in summer than in the winter (Figs. 24, 25 and 26). The map showing the 20°C winter isotherm with a temperature drop of 3°C (Fig, 25) indicates a tropical zone of considerable size and the possibility of a major faunal die-off i s improbable. If Briggs is correct, one would expect low affinities with the Indo-Pacific, which is the case, and perhaps closer affinities with the Atlantic due to intermingling in either the Sangaman interglacial period or late Pleiocene, Also, due to the lack of habitat, a poor east Pacific representation of coral reef fishes would be expected. Caranx distribution tends to support this idea. Hubbs* theory of a temperature drop in the neighbourhood of 3 to 4°C in the tropical east Pacific i s supported by most of the literature and is not argued here; however, i t is unlikely that such a drop had a serious effect on the shore fish fauna. The paucity of the shore fish fauna in this region is better explained by the barriers and lack of habitat. SUMMARY There are seven distinct species of Caranx in the tropical east Pacific area. These are: Caranx caballus, _C, vinctus. C. marginatus. C_. hippos, C_, lugubris. C. melampygus and C. medusicola. C, marginatus i s separable from C_« latus and C. sexfasciatus on vertebral counts, C. marginatus having one more caudal vertebrae. C. caballus L L H may be separated from C. crysos with a character index + — + ^ ) • C_. melampygus may be separated from C_, medusicola by the anal f i n shape and dimensions (C. melampygus - falcate portion pointed and long; C_. medusicola - falcate portion rounded and shorter) and snout shape (concave in CJ. melampygus; almost straight in C. medusicola). C. stellatus is synonymous with C_. melampygus; the previously reported colour differences being size dependent. The present distribution of east Pacific Caranx species may be described as: 2 - circumtropical (C. hippos, £. lugubris) 1 - Pan-Pacific (C. melampygus) 4 - endemic to the tropical east Pacific (C. caballus, C. marginatus, C, vinctus, C. medusicola) The present barriers to Caranx distribution in the tropical east Pacific are: temperature to the north and south; the Panama isthmus to the east; and the east Pacific sea barrier to the west. Non-endemic Caranx forms in the tropical east Pacific show different amounts of variability over their total range. C. marginatus : latus : sexfasciatus and C. caballus : crysos vary most between geographic regions and £. melampygus and £. lugubris vary least between geographic regions. The most probable reasons for the paucity of the tropical east Pacific - 55 -fauna are the presence of barriers, especially the east Pacific sea barrier, and the lack of coral reef habitat. Based on Caranx evidence i t is unlikely that the postulated 3 to 4C° Pleistocene drop in temperature had a serious effect on the shore fish fauna. BIBLIOGRAPHY Agassiz, L. 1829. In Spix, J.B. von. and L. Agassiz. Selecta genera et species piscium quos in itinere per Brasiliam annis 1817-20 Monachii. Pt. 1, 82 p., 48 p i . Arrhenius, G. 1961, Geological record on the ocean floor. Pp. 129-148 In Oceanography, Sears, M. ed. (1959) 1961. Publ. No. 67, A.A.A.S., Wash., D.C. Bartholomew, J. 1956. 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Ser. 13, Vi: 369-430. - 63 -- 6 4 -arangidae from the Tropical East Pacific "amily Carangidae Genus Selar Selar crumenophthalmus Genus Trachurus Trachurus picturatus Genus Hemicaranx Hemicaranx atrimanus Hemicaranx zelotes Hemicaranx leucurus Genus Caranx Caranx caballus Caranx vinctus Caranx melampygus Caranx medusicola Caranx lugubris  Caranx hippos Caranx marginatus Caranx sp. Genus Gnathanodon Gnathanodon speciosus Genus Citula Citula dorsalis Genus Alectis Alectis ciliaris Genus Vomer Vomer declivifrons Genus Chloroscombrus Chloroscombrus orqueta *65 Carangidae from the Tropical East Pacific (Cont.) Genus Selene Selene oerstedii Selene brevoortii Genus Trachinotus Trachinotus kennedyi Trachinotus culveri Trachinotus rhodopus Trachinotus paloma Genus Oligoplites Oligoplites altus Oligoplites mundus Oligoplites saurus Oligoplites refulgens Genus Elagatis Elagatis bipinnulatus Genus Seriola Seriola mazatlana Seriola dorsalis Genus Naucrates Naucrates ductor Genus Carangoides Carangoides orthogrammus A p p e n d i x 2 - C . cabal lus data ( A l l m e a s u r e m e n t s in m i l l i m e t r e s ) H e a d P e c t o r a l U p p e r E y e D o r s a l A n a l P e c t - C a u d a l G i l l L e n g t h D e p t h L e n g t h F i n Jaw L e n g t h F i n F i n o r a l D M V R a k e r s V e r t e b r a e Scutes L e n g t h L e n g t h - F i n m m 191.0 51 .3 50. 0 6 1 . 4 19.4 12.7 V H I - I 24 II--118 - - 7 21 8 28 10+ 15 44 185. 5 4 9 . 5 4 7 . 6 53 .7 18.0 12.0 23 19 8 19 8 30 10+15 43 186. 3 50 .2 47 . 8 57. 1 18.0 11.6 22 19 - - 30 - 42 271 .0 7 0 . 9 75 .6 - 29 . 5 - 22 19 - - 28? - 44 176. 1 4 6 . 9 4 6 . 2 50 .8 16.7 12.0 22 18 - - 27 ? 10+ 15 45 284. 1 7 4 . 2 7 5 . 2 97 .8 2 8 . 8 17.8 24 19 - - 30 - 45 203. 0 53 .3 52 .7 62 .0 19.6 - 23 20 - - 29 10+15 47 197.6 54 .6 51. 5 6 6 . 5 20. 1 - 24 19 8 20 7 30 10+ 15 46 196. 9 4 9 . 6 52 .0 58.6 19.0 - 23 20 8 20 8 28 10+ 15 49 280. 1 6 7 . 8 71 .2 92 .0 26. 5 - 24 18 - - 29 - 39 264 .7 6 9 . 0 6 9 . 9 8 5 . 8 26. 1 - 23 17 1-22 - - 28 - 41 167. 1 4 9 . 5 4 4 . 4 50 .5 17. 5 11.2 22 ' 20 - - 28 10+15 43 190.6 54 .4 52.7 59 .3 18 .8 11.6 24 20 - - 29 10+15 43 287 .7 7 5 . 7 73 . 5 98 .7 2 7 . 9 - 22? 20 1-23 - - 26? - 43 204 .7 57 .2 54 .0 - 19.6 - 23 19 - - 28 - 42 2 1 1 . 8 57 .2 54.7 6 3 . 2 2 0 . 8 - 23 19 1-22 - - 28 - 42 226 .6 64 . 1 58 .0 6 7 . 4 21 .6 - 24 20 - - 30 - 43 2 4 1 . 2 6 5 . 9 60 . 1 7 2 . 4 22. 5 - 22 19 1-22 - - 29 - 46 272. 5 7 6 . 0 72 . 5 9 2 . 8 2 7 . 8 - 23 19 - 28 - -326. 5 86 .7 85 .0 108.8 32. 1 - 22 18 - - 29, - 46 3 5 0 . 9 85 .0 9 4 . 5 127.4 36 .0 - 24 20 - - 30 - -199.6 4 5 . 7 51 .3 6 0 . 3 19.4 12.0 21 18 8 20 7 29 10+ 15 44 310 .6 6 4 . 9 78. 3 87 .0 2 9 . 2 - 24 19 - - 29 - 43 293 .7 7 7 . 0 82 .0 9 9 . 5 3 0 . 4 - 23 18 - - 29 - 46 168 .9 4 6 . 7 4 5 . 9 50 .9 17 .4 - 23 ? 16? - - - 25 - 45 283. 8 6 9 . 6 7 5 . 0 87 .7 2 7 . 9 16.4 24 20 - - 30 - 44 262. 0 6 7 . 0 •67.6 85 .7 2 5 . 9 17.0 22 21 - - 29 10+ 15 45 195.4 4 8 . 8 50.7 63 . 1 18. 6 13.0 23 19 8 19 8 28 10+15 44 Appendix 2 - (2) Length Depth Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gil l Length Fin Jaw Length Fin Fin oral D m M V m Rakers Vertebrae Scute Length Length Fin 173.0 47.3 45.4 52.6 16.7 11.4 22 19 _ 7 19 8 28 10+ 15 38 174.7 46. 1 46.9 50.4 17.5 11.5 24 21 I- 21 - - - 28 - 42 180.6 48.3 48. 9 56.6 18.4 12.3 24 21 - - - - 30 - 40 288. 80.4 75.2 92.6 28.0 16.9 22 19 - - - - 27 - 44 316. 84.0 85.0 112.3 33.2 16.9 23 20 - - - - 31 - 41 403. 95.5 106. 5 135.2 39.4 21.4 22 19 - - - - 30 - 42 393 95.8 105.6 126.6 39.4 23.2 21 19 - - - - 29 - 40 195.0 52.9 50.9 58.8 19.0 12.2 23 19 - - - - 29 10+ 15 49 310. 83.4 88.9 114.6 34.2 23.7 - 21 - - - - 28 47 197. 5 52. 5 52. 5 63.8 20.0 14.0 23 19 - - - - 28 - 41 180. 5 51.2 47. 1 56.9 16.8 10.8 23 19 - - - - 28 - 45 181.9 48.2 47. 1 55.7 18.4 11.7 23 19 - - - - 27 - 45 250. 64.0 67.9 81.0 25. 1 15.9 24 20 - - - - 27? - 41 ? 268. 71.9 73. 8 94.0 27.8 19.0 22 19 - - - - 29 - 43 314. 79.9 85.0 104.7 32.8 19.5 23 19 - - - - 29 - 46 343. 88.3 96.4 121.4 36.3 21.5 22 19 - - - - 27 - 45 322. 89. 1 90.4 115.4 35. 5 21.0 22 18 - - - - 30 - 42 297. 74.3 80.0 100.6 30.7 19.9 23 20 - - - - 28 - 44 313. 83. 1 84. 3 120.4 34.2 20.3 22 18 - - - - 31 - 41 300. 82. 1 81.8 103.7 29.6 17.0 23 20 - - - - 28 - 46 327. 83.2 85.8 109.4 32.9 18.9 20? 19 - - - - 30 - 40 314. 80.9 85. 1 105.2 31.6 18.8 22 18 - - - - 27 - 44 316. 83.2 83.6 106.5 32.0 21.0 23 19 - - - - 28? - 45 194.0 55.0 52.4 68.9 19.5 11. 1 24 19 I- 21 - - - 27 10+ 15 46 207.0 56.0 57. 5 64.1 20.6 12.0 23 19 I- 22 - - - 29 10+ 15 44 179. 1 49.7 46.7 50.2 17. 1 10.6 22 18 I- 22 - - - 27 10+15 44 203.0 61.2 54.8 69.9 21 .0 11.9 22 19 I- 22 - - - 30 10+ 15 51 212.0 56.8 56. 1 65.9 20.9 11.7 23 19 I- 23 8 19 7 29 10+ 15 51 220. 62. 1 58.2 74.6 22.0 12.9 23 - I- 23 9 17 8 31 10+15 47 Appendix 2 - (3) Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gill Length Depth Length Fin Jaw Length Fin Fin oral m M m Rakers Vertebrae Scutes Length Length Fin 218. 60.3 57.6 - 21.0 12.5 23 19 9 17 9 28 10+15 51 212 56. 5 57.2 69.3 21.4 12.9 22 18 1-22 - 29 10+ 15 43 294. 81.2 79.0 98.2 29.0 16.3 23 19 1-21 - 29 10+ 15 47 285 78.0 78. 5 100. 1 29.0 16.9 24 19 1-23 10+15 45 Append ix 3 - C . crysos data 184.2 55.0 51.5 60.3 21.7 12.0 VIII-I 24" II--I 19 1-22 - 26 10+ 15 48 177. 5 56.7 49.0 55. 1 19.4 12.2 . 21 20 1-21 - 27 10+ 15 46 175.3 54. 5 47.3 52.3 21.1 11.9 22 19 1-20 - 27 10+ 15 47 149.2 45.7 41.9 42. 1 16.9 9.4 24 20 1-24 - 26 10+ 15 45 145. 5 43.8 41.4 46.0 16.9 10.6 23 20 1-20 - 28 10+ 15 46 99.6 39.4 27.0 24.2 11.2 7.5 23 19 1-22 27 - 45 155. 5 48.4 41.5 41. 1 17.7 11.6 23 19 1-21 - 27 - 51 90.8 26.0 24. 8 17.3 10.3 6.5 24 19 1-20? - - 27 - 46 99.5 31.0 28. 2 23.6 11.4 8.1 23 19 1-21 - 27 - 48 165. 5 51.0 46.6 50.9 18.8 11.8 24 20 _ 26 10+ 15 45 Appendix 4 - C. vinctus data Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gill Length Depth Length Fin Jaw Length Fin Fin oral m M m ^ - a ^ e r s Vertebrae Scutes Length Length Fin 196.9 65.9 53.2 65.0 19.3 13.2 VIII-I 25 II-•I 21 - - 29 10+ 14 49 264.2 83.6 71.4 85. 5 26.7 - 22 21 - - 26 10+ 14 51 214.9 69.3 59.4 67.8 22.3 14.5 23 20 - - 29 - 51 253. 5 80.7 66.8 82.2 24.4 18.4 23 19 - - 29 10+ 14 52 172.7 54.8 45.2 57.0 16.8 11.1 24 20 - 8 18 10 28 10+ 14 51 167.8 53.0 44.6 50.7 16.6 11.1 24 21 - 9 17 8 28 10+ 14 49 92.2 29.5 26. 1 25.1 ^ 9.2 6.5 24 19 1-18 9 19 9 30 10+ 14 48 102.8 33.2 28.4 28.5 9.8 7.3 23 20 1-18 10 18 10 29 10+ 14 48 100.9 32. 5 28.9 28. 5 10.0 7. 1 23 20 1-19 10 18 10 28 1P+ 14 47 284.0 85.2 73.9 - 82.9 27.0 15.5 23 19 1-18 10 18 9 29 10+ 14 52 118.7 41. 1 31.9 34.6 11.8 8. 1 23 20 1-20 - 30 10+ 14 50 130.8 42.2 35. 8 38.7 13.0 8.7 23 20 1-19 - 27 10+ 14 45 126.7 41.0 34. 5 39. 1 12.2 8.8 24? 20 1-18 - 30 10+ 14 47 127. 2 42.0 34. 1 37.8 12. 1 9.1 24 20 1-18 - 28 10+ 14 47 126.2 40.0 32.9 37.7 12.2 9.0 23 20 1-18 9 18 -9 28 10+ 14 51 124. 8 40.0 32.6 36.8 11.3 8.8 23 20 1-20 11 18 9 28 10+ 14 48 161.0 56.9 44.6 47.9 14.2 8.8 23 20 1-19 - 27 10+ 14 51 59.9 22.2 16. 3 12.0 6.1 4.6 22? 20 1-19 - 28 10+ 14 47 239. 78.9 64.6 79.9 23.0 13.8 22 20 1-19 - 28 10+ 14 48 266. 85. 1 72.2 83.0 27.0 15.2 24 20 1-19 - 29 10+ 14 48 263. 86. 3 70.8 79.5 26.3 15.1 23 19 1-19 - 29 10+ 14 50 231. 74.2 61.3 75.6 22.6 13.9 23 19 1-19 - - 10+ 14 51 Appendix 5 - C. hippos data Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gill Length Depth Length Fin Jaw Length Fin Fin oral D M V Rakers Vertebrae Scutes Length Length Fin 246. 79.9 74.8 87. 1 32.9 15. 1 - - r-22 - 17 - -243. 82. 1 74.2 84.0 33.3 15.2 VIII -I 20 II-I 16 21 - 18? - 39 262. 85.2 78.0 86.6 33.8 16.2 19 16 20 - 17 - 37 289. 95.6 87.7 101. 8 39. 1 15. 5 19 17 20 - 18 - 41 215. 80.0 64.2 72.2 28.2 14.4 18 16 19 - 17 - 39 219. 74.3 66.2 70.0 28.9 14.3 20 17 20 - 17 10+ 14 44 234. 86.4 69.9 76.2 30.0 15. 1 20 16 20 - 18 - 42 215. 75.5 66.6 73. 1 27.9 14.3 20 16 20 9 19 8 17 10+ 14 40 202.0 71.2 61. 5 69.4 27. 1 14.2 20 16 20 8 18 9 17 10+ 14 39 185.4 63.8 55.0 61.5 24.2 13.0 19 16 20 - 18 - 37 163.4 56.4 48.6 52.6 22. 1 12.6 19 16 20 - 18 - 38 186.3 65.6 55. 8 61 .0 24.3 13.3 20 17 21 - 17 10+ 14 41 172. 1 60.3 52.7 56.7 22.2 12.8 19 16 20 - 18 10+ 14 40 168.7 59.9 50.9 55.6 22.0 12.4 19 16 20 - 18 - 41 167. 8 60.0 51.0 52. 5 22.0 11.3 19 - 20 9 18 9 18 10+ 14 37 145.7 52.9 43.4 47.2 19.3 10.9 19 16 19 8 18 8 18 10+ 14 38 67.0 25.6 20.2 17.8 8.5 - 20 17 - - 19 - 35 111.0 39.3 31; 8 28.2 13;6 - 20 18 - - 18 10+ 14 36 107.8 39.1 31.2 31.0 13.3 - 21 17 - - 16 10+ 14 35 93.0 34.4 26. 6 24. 5 11.9 - 21 17 - - 17 10+ 14 35 68. 5 25.5 19.9 13. 5 8.4 - 20 17 - - 18 - 32 345. U 5 : 100; 120. 45.0 - ? 20 16 - - 16 - 36 380. 130. 115. 135. 50. - •? 20 17 - - 15 - 38 550. 160. 160. 175. 70. - ? 21 16' - 16 - 37 110.0 39.3 33.2 33.5 14.4 - VIII-I 21 17 - 8 17 8 16 10+ 14 38 113.6 40.9 33.0 35. 1 15. 3 - 20 17 - 8 19 8 17 10+ 14 34 172.2 61.2 51.0 54.0 23. 1 - 20 17 - - 17 - 37 171.6 62.4 51. 8 55.6 23. 1 - 21 16 - - 17 10+ 14 37 108.0 43.8 34.6 35.2 15.6 . - 21 16 - - 16 - 36 124.6 45.0 32.0 35.4 14. 5 - 20 16 - - 17 - 37 Appendix 5 - (2) Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gill Length Depth Length Fin Jaw Length Fin Fin oral m M V Rakers Vertebrae Scutes Length Length Fin 115. .4; 40.4 33.9 32. 3 14.0 - VIII--I 21 II-I 16 - - 17 36 143.0 52.2 41.8 40.7 17.9 - 21 16 - 16 _ 37 322.9 96.9 95.0 106.0 41.0 - 21 17 - 15 _ 40 263.9 76.6 73.7 85.2 32.8 - 22 17 - - 16 38 150.8 52.2 45.6 49.2 20.8 - 20 16 - - 15 10+ 14 35 108.7 38.8 32.2 31.4 14.7 - 21 16 - - 16 - 32 127. 1 47.0 37. 1 37.7 16.7 - 20 17 - 16 _ 37 123. 1 43.4 39.5 37.6 16.5 - 21 17 - 16 _ 34 355.0 110.7 102. 5 112. 1 44.9 - VII-•I 20 16 - — 15 _ 36 243. 5 89. 1 73. 3 86.8 33.7 - VIII -123 16 - - 16 10+ 14 35 219.4 74.9 66.4 73. 1 28.4 - 20 16 - 8 19 9 15 10+ 14 34 178. 1 63. 1 52. 5 55.7 22.9 - 20 16 - - 15 :: t. ••• 39 163.0 57.8 47.3 48. 5 20.3 - 20 16 - - 16 - 38 182.3 64.8 54.8 58.6 24.4 - 20 16 - - 16 - 40 112.6 38.5 34.0 33.4 14.3 9.9 20 16 - - 17 - 28 104.0 37. 5 31. 1 .29.7 13.0 9.5 21 15 - - 18 - 29 96. 1 32.5 27.4 29.4 11.9 8. 5 21 16 - - 17 28 66.4 24.2 20.2 17.4 8.8 6.0 20 ? 16? - - 18 - 27 123.0 42.1 35. 1 33. 5 / 15.3 19.4 10.2 23 17 - - 17 _ 33 142.9 49.6 43. 1 45.0 11.4 20 16 - 17 _ 35 168.4 57.9 50. 8 53.0 22.6 13.6 20 16 - - 17 10+ 14 41 68.2 25.9 19. 1 16.3 8.4 5.4 20 16 - - 17 10+14 33 198.0 63.9 58. 1 66. 1 25.5 14.3 20 16 - - 16 10+14 39 196.0 66.3 59.6 64.6 26.7 13.9 20 16 - - 16 10+ 14 37 444. 125. 133. 154. 60. 23. 20 20 16 17 1-22 8 18 8 16 10+ 14 10+ 14 -- - - - - - 20 17 20 8 18 7 - 10+14 _ 508. 155. 147. 168. 63. 25. 21 17 - - 16 - 34 486. 149. 147. 176. 62. 24. 19 16 - - 16 - 38 730. 195. 210. 237. 89. 29. 18 17 - - 16 - 37 Appendix 6 - C. marginatus' data Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gill Fin Jaw Length Fin Fin oral Length Length Fin Length Depth Length  D^ M V^ Rakers Vertebrae Scutes 186.-0 65.8 52.8 54.9 25.9 14.3 VIII-I 21II-I 16 - - 16 - 31 243.3 81.0 69.3 80. 1 32. 1 17. 1 21 16 - - 16 - 31 241.2 78.8 69.9 82.7 31.9 16.3 21 16 - - 15 - 33 91.0 33.2 28. 1 25.5 12.7 7.3 20 15 1-22 - 16 10+15 28 101. 1 33.7 28.9 25.8 13.3 8. 5 20 15 20 - 16 10+ 15 30 100. 1 34.3 30.7 28. 5 14.3 8.2 20 17 22 - - 10+15 31 87.4 30.9 25.0 21.8 11.0 7.0 19 15 20 - 16 - 31 116.0 41.7 33. 1 30.7 15. 1 8.2 21 17 20 - 16 10+ 15 33 93.6 33.7 27.4 23.2 12. 1 7.7 20 15 20 - 16 10+15 29 97.4 35.0 27.0 23.2 11.5 7.6 19 15 21 - 17 10+ 15 29 90.7 31.1 26. 5 23.7 11.9 8.0 21 16 21 - 16 10+ 15 31 89.6 31 .8 24.6 20.3 11.5 7. 1 21 16 21 - 17 10+ 15 30 86.2 30.5 25.9 20. 1 10.3 6.6 20 16 21 - 17 10+ 15 30 127.7 46.8 38.0 38.6 16.9 10.1 20 15 21 - 17 10+15 30 122.9 44.1 36.8 34.7 17.2 9.8 19 17 21 8 17 9 17 10+ 15 30 122.4 43. 5 34.8 34.7 15.6 9.7 19 16 20 9 18 8 17 10+15 32 118.2 41.9 34. 5 32.4 15. 1 10.0 20 17 18 7 19 7 17 10+ 15 31 124.3 44.9 36.4 35.3 15.3 9.9 19 14 20 8 18 7 16 10+ 15 31 118.0 42.7 35.8 35.8 15.3 10. 5 20 15 19 - 17 10+ 15 30 125.0 44.3 36.6 36.6 16.5 9.9 20 15 19 - 17 10+ 15 28 113.6 39.8 33.4 32.2 14.8 9.1 19 16 20 - 17 10+ 15 30 110. 8 39.3 32.7 32.8 14.8 8.8 19 15 20 - - 10+ 15 31 128. 1 44.8 36.7 38.3 16.8 10.0 21 16 20 - - 33 113. 1 40.3 34.0 34. 1 15.8 9.9 20 16 20 - - 31 104. 5 36.7 30.0 27.8 13.8 - 20 17 - - 16 10+ 15 30 108.6 38.2 30.7 30.0 14.3 8.9 19 16 20 - 17 - 3.1 140.0 49.4 40.0 40.0 18.8 11.0 20 16 21 8 17 9 17 10+15 31 140. 8 46.1 40.6 41.9 19.1 11.5 20 16 21 8 17 9 17 10+ 15 36 Appendix 6 - (2) Head Pectoral Upper Eye Dorsal Anal Pect- Caudal G i l l Length Depth Length ,Pin,, Jaw Length F i n Fin oral D M V Bakers Verte- Scutes e n ® Length Fin brae 127.5 42.4 36.8 37.5 18.0 11.0 VIII-I 21II-I 17 I- 22 — 16 10+15 30 119.5 40.1 36.2 37.4 16.8 9.8 19 15 22 — 17 10+15 28 137.8 44.4 38.4 39.0 18.0 10.3 20 16 21 — 17 — 30 118.0 41.2 35.1 36.4 16.7 10.0 20 16 21 — 18 — 29 175.6 60.9 54.3 56.7 25.5 14.5 20 16 — — 16 10+15 31 147.6 48.7 43.5 45.5 20.6 13.0 20 17 — — 15 10+15 29 152.2 49.2 43.6 46.0 20.3 13.0 20 16 — — 15 10+15 27 116.0 42.0 35.4 33.8 16.0 9.8 21 17 — — 17 10+15 33 106.0 38.6 29.2 — 13.5 7.8 20 17 — — 16 10+15 33 221.3 75.9 68.9 64.4 29.3 17.9 20 16 — — 16 — 34 201.0 67.9 57.3 61.4 26.8 15.4 20 16 — — 15 10+15 29 191.3 68.1 55.4 58.7 25.1 13.3 21 17 — — 16 — 30 110.5 36.7 30.4 29.8 13.5 7.5 20 16 — — 16 10+15 29 100.1 33.5 28.8 29.2 12.7 8.6 20 16 — — — 10+15 31 127.7 45.0 36.4 33.4 16.1 9.6 21 16 — — 16 — 32 148.0 59.1 43.1 41.9 19.3 12.2 20 14 — — 15 10+15 31 175.0 61.1 50.0 57.0 23.6 15.5 19 16 — — 15 10+15 31 170.0 58.1 49.7 53.4 23.1 14.0 21 16 — — — 10+15 28 152.8 52.1 46.0 46.2 20.2 11.7 21 16 — — 16 10+15 30 153.0 50.9 45.7 45.1 20.9 11.6 21 16 — 9 18 8 16 10+15 30 123.1 43.5 35.3 34.9 16.5 10.9 21 16 — 8 17 8 16 10+15 34 177.0 61.8 53.0 55.4 24.6 13.6 21 17 20 17 10+15 36 191.2 67.4 57.9 60.9 26.4 15.1 19 16 20 — 17 10+15 32 202.1 69.0 60.4 64.8 28.3 16.3 21 17 20 — 17 10+15 35 211.0 73.5 63.0 67.9 29.0 16.4 20 17 20 — 17 10+15 32 213.0 76.1 66.2 71.1 30.8 17.4 20 17 — - 17 — 31 170.1 60.3 52.0 54.5 24.1 14.3 20 15 19 — 16 — 34 177.9 62.7 52.5 59.4 23.7 14.0 18 15 20 — 17 10+15 31 164.2 56.2 50.9 50.2 23.2 12.5 20 ,16 20 — 16 10+15 32 168.1 50.8 49.6 49.0 23.3 13.8 20 16 21 — 16 - 34 157.3 56.1 48.3 48.5 21.4 13.5 20 16 21 — 17 - 30 Appendix 6 - (3) Head Pectoral Upper Eye Dorsal Anal Pect- Caudal G i l l Length Depth Length Fin Jaw Length Fin F i n oral D M V Rakers Verte- Scutes Length Length Fin brae 150.0 55.9 49.4 50.8 21.6 12.8 VIII 173.1 61.2 52.6 — 23.7 14.7 153.0 54.1 45.3 48.0 21.3 13.1 151.9 23.7 47.1 48.5 20.6 12.8 149 .9 53.6 44.1 46.8 20.0 12.4 Appendix 7 - C. latus data 160.0 60.0 48.4 48.0 23 .9 13.0 V I I I 112.0 39.5 34.1 29.3 14.8 9.4 87.9 32.0 26.8 24.8 12.9 8.4 53.2 20.1 19 .9 12.6 7.5 5.4 85.0 30.9 26.9 22.2 10 .9 7.0 I 20 11-116 22 8 18 8 16 10+15 33 20 16 22 7 19 9 16 10+15 33 20 17 20 — 18 10+15 34 20 16 22 — 16 10+15 31 20 15 21 — 17 10+15 32 ] I 20 II-I 18 20 8 17 7 17 10+14 36 20 17 19 9 16 8 17 10+14 38 21 16 20 8 17 8 16 10+14 35 20 17 — 3 — i 16 10+14 33 21 16 21 \% 7 16 10+14 33 \ Appendix 8 - C. lugubris data Length Depth Head Length Pectoral Pin Length Upper Jaw Length Eye Dorsal Length Pin Anal Pin Pect-oral Fin Caudal D M V ra ra G i l l Rakers Verte-brae Scutes 228.9 93.5 65.1 77.3 26.3 14.0 VIII-I 22 II-I 17 _ 19 _ 30 287.6 108.9 86.9 99.5 36.0 21.5 20 18 — — 20 — 27 310.3 107.2 90.0 107.2 37.0 22.1 22 18 — 9 17 8 17 10+14 28 204.4 73.3 60.0 72.3 22.6 13.4 22 19 — 8 19 8 18 10+14 30 197.1 67.1 57.5 63.5 22.7 13.5 21 17 - 8 20 8 19 10+14 32 755. 252. 220. 230. 91.5 36. 21 18 — — — — 34 372. 123. 112. 117. 45.4 27.2 21 17 r-21 — 19 10+14 27 354. 126. 107. 122. 42.2 26.1 21 17 21 — 18 10+14 29 493. 168. 144. 162. 58. 32. 20 17 22 - — — 28 566. 187. 158. 174. 65. 26, 20 17 21 - — — 27 393. 134. 114. 137. 47. 24. VIII-I 21 17 22 - — - 27 392. 126. 111. 123. 47. 24. 21 18 22 — — — 31 450. 148. 131. 143. 52. 29. 21 17 21 - — — 29 355. 122. 110. 126. 42. 26. 21 18 21 — — — 29 421. 141. 124. 142. 49. 26. VII-I21 17 23 — - - 28 349. 127. 109. 121. 42. 24. VIII-I 22 19 22 — — — 27 579. 188. 164. 188. 66. 32. 22 18 22 — — — 28 466. 157. 137. 154. 54. 31. 21 17 21 — - - 30 400. 140. 119. 137. 47. 28. 21 18 20 — — 28 621. 215. 176. 180. 71. 35. 20 19 23 — - - 30 365. 131. 110. 129. 43. 25. VIII-I 21 16 22 - - - 32 Appendix 9 - melampygus data Head Pectoral Upper Eye Dorsal Anal Pect- Caudal G i l l Verte-Length Depth Length Pin Jaw Length - Pin Pin oral D M m V m Rakers brae Scutes Length Length Pin iii129.2 51.4 37.4 40.2 15.1 9.9 VIII-I 21 II-I 19 1-21 9 16 9 18 10+14 35 109.3 41.7 32.2 34.0 12.6 8.5 22 19 21 9 17 9 19 10+14 35 104.4 39.1 32.4 33.2 12.0 8.9 21 18 21 9 17 8 19 10+14 33 89.0 36.1 29.0 27.0 10.0 7.1 21 18 21 9 17 8 19 10+14 34 241.2 83.3 67.1 78.4 26.9 13.9 22 19 22 — 19 — 34 214.4 78.5 61.1 65.4 23.5 13.1 23 20 20 — 19 10+14 36 209.9 76.9 57.6 69.9 23.1 12.9 22 19 21 — 19 10+14 35 234.9 86.9 65.1 83.9 25.6 13.3 22 20 22 — 19 10+14 36 151.0 57.7 43.0 48.5 16.9 10.0 23 19 21 — 18 10+14 33 106.6 40.8 30.8 32.6 11.9 8.2 22 20 22 — 19 10+14 37 118.5 44.5 33.9 13.8 13.8 9.4 23 19 22 — — 10+14 35 , 398. 136. 124. 128. 49. 22. 22 19 20 — 19 — 29 ^ 32 ? 111.4 43.8 31.9 35.0 12.6 9.6 VIII-I 22 20 — — 19 10+14 131.2 52.8 35.4 42.6 14.8 8.9 22 18 — — 19 10+14 33-4 115.1 43.2 32.5 38.1 13.0 9.3 22 19 — — 19 10+14 33 118.1 45.0 34.7 37.7 13.7 8.9 22 19 — — 19 - 33 142.8 52.2 41.8 49.9 16.5 10.6 22 19 — — 18 10+14 35 107.2 40.1 30.5 34.5 12.3 8.3 23 19 — — 19 10+14 34 121.5 43.8 35.1 40.7 13.0 9.0 22 18 — — 18 10+14 33 109.6 42.0 32.0 33.5 12.4 8.4 23 18 20 — 19 10+14 36 114.8 42.7 31.3 35.0 12.8 8.9 22 19 21 8 17 } 9 — 32 115.2 41.9 32.5 36.1 12.6 8.6 22 19 21 9 19 10+14 34 117.2 43.0 33.8 35.8 13.1 8.8 22 18 21 9 17 8 18 10+14 36 116.7 42.9 33.6 36.2 13.5 9.6 VII-I 22 19 22 8 19 8 19 10+14 37 118.6 46.0 32.9 36.0 13.2 9.7 VIII-I 22 18 22 8 19 7 19? 10+14 34 111.9 42.3 31.4 34.2 12.6 9.3 22 19 22 — 19? — 34 120.5 43.6 35.0 36.8 12.8 8.4 22 19 21 — 18 — 36 114.0 42.2 31.4 35.0 12.3 8.6 22 19 20 — 18- — 35 111.4 40.6 31.4 32.1 12.8 8.3 22 19 21 - 19 — 34 115.2 41.3 32.1 37.8 12.4 8.7 22 19 22 — 19 - 33 610. 203. 183. 197. 79.4 26.0 21 16 21 — 18 - 31 441. 153. 131. 143. 50. 25. VIII-I 21 18 19 — - — 31 636. 207. 190. 192. 80. 30. 21 18 22 - - - -Appendix 9 - (2) Head Pectoral Upper Eye Dorsal Anal Pect— Caudal G i l l Verte-Length Depth Length Pin Jaw Length Fin Fin oral D M V Rakers brae Scutes Length Length Fin m m 654. 247. 204. 224. 83. 26. VIII-I 22 II-I 29 — — 34 642. 213. 199. 240. 82. 28. 22 20 — — 17 _ 33 672. 223. 201. 230. 83. 28. 23 19 — _ 16 _ 34 670. 210. 197. 225. 81. 28. VII-I 22 18 — 17 — 33 Appendix 10 - C. medusicola data 113.9 44.0 31.5 30.0 11.9 7.3 VIII-I 22 II-I 20 r-21 19 10+14 37 111.7 44.3 30.6 30.2 12.0 7.1 23 20 21 — 19 10+14 37 157.2 64.2 43.9 30.2 18.1 10.4 21 19 21 — 19 10+14 40 

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