A REVIEW OF THE 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 partial fulfilment of the requirements for an advanced degree at the University of

British Columbia, I agree that the Library shall make it 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 is 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 it 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 critical 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. - iii -

TABLE OF CONTENTS Page

ABSTRACT i ACKNOWLEDGEMEM: ii TABLE OF CONTENTS iii LIST OF TABLES vi LIST OF FIGURES vii

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 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 Gill 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 - vi -

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 - vii -

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 fin 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 al, 1942) . -. 48b INTRODUCTION

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. MATERIALS AND METHODS

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 is 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.

All 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 fin 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). Gill rakers were counted on the

lower portion of the first right hand gill arch, including all rudiments which

were sufficiently developed to be felt with a fine probe. Lateral scutes

counts include those specialized scales on the posterior half of the lateral

line (straight portion), starting at the first scale with a pointed posterior 4

3

1- Fork length 4-Head length 2- Depth 5- Eye length - 3- Pectoral fin length 6- Upper jaw length

Figure 1. Measurements made on Caranx species. 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 al, 1953) will not be used here. By these definitions, in Caranx there is one short spine on the dorsal edge of the pectoral fin. All counts, measurements and characters used are based on an examination of all 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.

No* of Specimens of each Character Species Examined

Relationship of ventral fin 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 little within the genus Caranx. Fig. 2 is an X-ray photograph of C. hippos. Abdominal vertebrae (those in front of the anastomosed - 3a -

Fig. 2 X-Ray photograph of C. hippos 1. Dorsal pterygiophore spine 2. First abdominal vertebra 3. First caudal vertebra 4. Last caudal vertebra 5. First major caudal ray 5'. Last major caudal ray

- 4 -

first 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 all species. Between any two species if 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 it 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 fin length or any proportion involving pectoral fin length was found to be of little value as the very long, narrow trailing end of the fin is subject to breakage and it is often impossible to ascertain if 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 difficult 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,

i i « range a* 2 standard deviations about ®ean (J) • a a G. caballus 167-241 mm 31 specimens d = C. marginatus 86-243 ©a 64 specimens a'a c. caballus 250-403 mm 30 specimens e =* CT lugubris T97-220 so 3 specimens b n 57 yinctua 60-284 mm 22 specicseno e'* C7 luiiubria 288-755 mm 25 specimens c a C. hippos 66-246 m 46 specimens f a C« aelampygua 89-241 on 29 specimens c' = C. hippos 262-730 mm 12 specisens f'n raelaapymia 398-675 as 8 specimens

g = C. aedualcola 112-157 ®m 3 epecioens - 4b -

Figure 4. Length/depth, aymbola ae in Figure 5* 2.0 2.5 3.0 3.5 4.0 4.5 5.0 5.5 6.0 —i— —i— —i— _H E

-1.93 •70S-

P

Figure 5. Head length/eye length and length/pectoral fin length symbols as in Figure 3 except a = caballus (all specimens) b = £i vinctus (all specimens) c = 9JL hippos (all specimens) d = marginatus (all specimens) e = lugubris (all specimens) f = C_j_ melampygus (all specimens) g = C. medusicola (all specimens) - 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; Is. = 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 titles of the following nature were encountered, "A

Collection of Marine Pish from the Pacific Coast of Mexico with some

Descriptions". All such titles were checked, many of which made no reference to Caranx. or perhaps only a distribution record. These titles are not included in the present bibliography. The titles 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 all 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 first 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 is 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 fin 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. Gill opening wide, gill membranes usually not united, free from isthmus. Branchiostegals 7. - 8 -

Gill arches 4, a slit behind the last. Scales complete or incomplete, cycloid, often deciduous. Dorsal fins two, the first spinous with 3-8 spines, moderate in height to very low. The pterygiophore bearing the first 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 fin with 1 spine followed by 14-30 rays and preceeded by two individual spines. Detached finlets occasionally present after dorsal and anal fins. Pectoral fin 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 all 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 difficult 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 it 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.

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 finlet. 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 it Selar

Gill Cover — Fleshy Projection

DD Shoulder girdle without furrow or fleshy projection. - 10 -

E Lateral line with bony scutes along its 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 difficult 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. Selene

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. Oligoplites

NN MaxiHilarm y not large - never less than 2 2/3 ini head ; dorsal and anal fins falcate. Scales not deeply embedded, skin not leather-like. Trachinotus

LL Anal fin much shorter than second dorsal, its base shorter than abdomen.

0 Dorsal and anal each with a single detached finlet. Elagatis

00 Dorsal and anal without finlets.

P First dorsal with 6-8 slender spines. Seriola

PP First dorsal with 3-4 low stiff spines. Naucrates

This character is uncertain, but the pectoral fin of Selene is much longer than that of either Oligoplites or Trachinotus.

This character is 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 first designated by Lacepede (1802) who apparently took the name from Commerson (pre Linnean), Jordan and Evermann (1896), present a history of Caranx which will 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 it to Caranx speciosus (now Gnathanodon). In printed nomenclature

Caranx was first used by Lacepede; who adopted the name from Commerson applying it 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 (Gill,

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. Gill 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 fin strongly forked, the peduncle slender, adipose ridges above and below lateral line half on peduncle and half on anterior caudal fin. Scales small, cycloid, often irregular and sometimes deciduous.

Breast scaled except in C. hippos where it 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 life.

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 it 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 Gill. 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 fig. 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

Size Range Cat. No. Specimens Locality (mm.)

W51-39 4 Tenados I., Sinaloa, Mexico 212-220 ¥51-49 1 Venados I., Sinaloa, Mexico 207 W51-51 2 Venados I., Sinaloa, Mexico 285-294 W51-56 1 Mazatlan, Sinaloa, Mexico 194 W59-12 1 Point Willard, Lower California, Mexico 203 W59-248 1 Pulmo, Lower California, Mexico 179 BC54-46 3 Acapulco, Guerrero, Mexico 186—191 BC54-52 1 Socorro I., Revillagigedo I *, Mexico 351 BC55-38 2 Zihautanejo, Guerrero, Mexico 175-181 BC56^337 1 Indefatigable I., Galapagos Is., Eq. 262 BC57-94 1 Petacalco Bay, Guerrero, Mexico 195 BC57-95 1 Petacalco Bay, Guerrero, Mexico 284 BC57-98 4 Petacalco Bay, Guerrero, Mexico 181-310 BC57-104 1 Santiago Bay, Colima, Mexico 343 BC57-108 1 Navidad Bay, Compilo, Mexico 200 BC57-144 2 Socorro I., Revillagigedo I ", Mexico 250-268 BC57-168 1 Maria Magdelena I., Tres Marias Is*, Mexico 271 - 16 -

BC59-236 1 Cape San Lucas, Lower California, Mexico 176 BC59-256 1 Clarion I., Revelligigedo Is*, Mexico 169 BC59-270 1 San Juanito I., Las Tres.Marias I *, Mexico 284 BC59-680 3 Chame Point, Panama 197-203 BC59-689 3 Panama City Mkt., Panama 205-212 BC60-5 2 Manjanillo Mkt., Colima, Mexico 294-311 BC60-15 1 Acapulco Mkt., Guerrero, Mexico 327 BC6CKL13 2 Tabago I., Panama 265-280 BG60-114 2 Tabago I., Panama 167-191 BC60-116 3 Tabago I., Panama 227-273 BC60-465 2 Isla la Granda, Guerrero, Mexico 173-195 BC60-466 2 Isla la Granda, Guerrero, Mexico 297-313 BC60-482 1 Maria Magdelena I., Las Tres Marias Is* ,Mex,,31 4 BC60-485 1 Maria Magdelena I., Las Tres Marias IS\Mex,,31 6 BC60-502 6 Cape San Lucas, Lower California, Mexico 195-403 BC61-142 1 Maria Magdelena I., Las Tres Marias I' Mex, ,314 BC61-181 1 Socorro I., Revellegigedo Is., Mexico 322

Description

Dorsal VIII-I, 21-24 (one specimen 20 but with damaged fin); 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. Gill rakers on lower limb of first 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 fin 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 fin. 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

BC60-465 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 fin base.

Teeth present on both jaws, vomer, palatine and tongue, jaw teeth deciduous in 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 life), ventrally pale yellowish (silver in life).

Distribution: (See Fig. 7)

C, caballus is endemic to the tropical East Pacific area. The southward limit is on the Peruvian Coast, probably Cape Aguja, (Walford, 1937). To the

North it 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 fig. 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 197 BC60-25 1 Port (Puerto) Arista, Oaxaco, Mexico 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 fin); 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. Gill rakers on lower limb of first 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 fin 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 fin. Rayed dorsal and anal fins with reduced scaly sheath on anterior portion of fin 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 life), ventrally pale yellowish (silver in life). 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 it 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. Caranx vinctus BC60-30 specimen No.1

Figure 8. Figure 9, - 20 - -

CARANX HIPPOS LINNAEUS (Pig. 10)

Synonymy

Scomber hippos Linnaeus, 1766, p. 492 (desc.)

Carangus chrysos Gill. 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 fig. 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. Is*) 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

Size Range Cat. No. Specimens Locality (mm.)

¥51-22 16 Mazatlan, Sinaloa, Mexico 146-289 BC56-142 1 Capoblanco, Peru 550 BC56-145 1 Capoblanco, Peru 444 BC57-78 2 Acapulco, Guerrero, Mexico 196-198 BC57-87 6 Petacalco Bay, Guerrero, Mexico 108-172 BC57-118 2 Chamela Bay, Colina, Mexico 345-380 - 21 -

BC59-215 5 Mazatlan, Sinaloa, Mexico 163-244 BC59-656 1 Cupica Bay to Orpu River, Columbia 69 BC59-667 4 30 M. N. of Panama City,-mouth Caimito River, Panama 113 BC59-671 3 Santa Clara, Panama 93-111 BC59-674 2 Los Angles, Panama BC59-676 1 Los Angeles, Panama 355 BC59-685 4 Chiman Area, 90 M. S. Panama City 109- 150 BC59-687 2 Panama City Mkt», Panama 264-323 BC60-12 2 Manzanilla, Coliraa, Mexico 68-168 BC60-25 2 Puerto Arista, Oaxaco, Mexico 110- 114 BC60-28 1 San Bias, Nayarit, Mexico 143 BC61-I26 1 Acapulco, Guerrero, Mexico 123 s N 906 1 Maria Madre?, Tres Marias I *, Mexico 508 N 907 1 Maria Madre, Tres Marias Is*, Mexico 468 N 708 1 Cape San Lucas, Lower California, Mexico 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. Gill rakes on lower limb of first arch 15-19 (usually 16-18).

Body moderately compressed, snout steep but rarely if 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 fin 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 fin 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 fin. 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 fin rays on posterior 3/4 of fin; caudal yellowish with dusky edges; anal yellowish with dusky markings on edge. Ventral fin yellowish; pectoral fin 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 life), ventrally pale yellowish (silver in life). Four or five vertical dark bars across body reaching to below the midline in young, disappearing in adults.

Distribution (See Fig. ll)

C. hippos is regarded by several authors to be circumtropical (Berry,

1959; Briggs, I960). However, several fauna works claiming to include all

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

BC57-87 specimen No.1

Figure 10. - 22b -

Figure 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 it 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 Gill. 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 Is* 140 BC56-445 1 Tower I., Galapagos Is* 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 Is', Mexico 148-176 BC59-202 2 Maria Magdalena I., Tres Marias ISJ Mexico 106-116 BC59-241 5 Maria Magdalena I., Tres Marias IsJ Mexico 123-175 BC59-264 1 Cleopha I., Tres Marias Is*, 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 first 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 fin 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 fin 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 fin 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 fin; anal pale yellowish; caudal dusky yellow with darker posterior border. Ventral fins pale yellowish. Pectoral fin 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 gill opening. Body dark dorsally (irridescent blue in life), ventrally pale yellowish (silver in life). 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»

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 Is* Caranx marginatus ; BC59-241 specimen - No. 2

Figure 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 al, 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 Is*, Mexico 288 BC54-403 2 Cocos I., Costa Rica 197-204 BC57-159 1 Socorro I., Revillagigedo Is', Mexico 310 BC58-383 1 Clarion I., Revillagigedo Is*, Mexico 229 BC61-179 13 Socorro I., Revillagigedo Is', Mexico 349-621 BC61-183 7 Socorro I., Revillagigedo Is*, Mexico 325-421 BC61-184 2 Socorro I., Revillagigedo Is', Mexico 354-372 BC61-187 1 Socorro I., Revillagigedo Is*, 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. Gill rakers on lower limb of first 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 fin 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 fin. Rayed dorsal and anal with very reduced scaly

sheath on anterior portion of fin only. Pectoral fins very long and falcate

in adults. Rayed dorsal and anal fins very falcate, longest rays of the fin

equal to more than l/2 the length of the fin 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 still 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 it 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, it is present around islands in both the Pacific and the Atlantic, but is not recorded to date from the mainland of America. C. lugubris is 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 fig. 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 al, 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

Size Range Cat. No. Specimens Locality (mm.)

rS. BC57-152 2 Socorro I.» Revillagigedo , Mexico 107-151 BC58-383 18 Clarion I., Revillagigedo , Mexico 110-131 BC59-261 4 Socorro Revillagigedo , Mexico 210-241 I., s. BC59-264 4 Cleopha I., Tres Marias I Mexico 89-129 BC61-149 3 Maria Madra I., Tres Marias Is *, Mexico 642-675 BC61-179 3 Socorro I., Revillagigedo I * Mexico 441-610 BC61-181 1 Socorro I., Revillagigedo I * Mexico 398 BC61-187 1 Socorro I., Revillagigedo I * Mexico 670 s BC61-189 1 Socorro I., Revillagigedo I * Mexico 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. Gill rakers on lower limb of first

arch 18-19 (rarely 16-17). Body compressed, snout fairly 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 fin 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 fin. Rayed dorsal and anal fins falcate, the longest rays equals from 2/5 to slightly more than l/2 of the length of the fin 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 all 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 life), ventrally silver as in life, 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 life), ventrally dirty silver as in life; 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 all fins a little 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 life), ventrally a golden shade as in life, many spots on side of body and head.

From the above it 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

it is recorded from the Tres Marias Is* 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 fig. 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. Size Range Cat. No. Specimens Locality (mm.)

BC54-64 2 Cape San Lucas, Baja California, Mexico 112-114 BC60-488 1 Maria Magdalena ILas Tres Marias Is*, 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. Gill rakers on lower limb of first arch 19. Body compressed, snout fairly 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 fin 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 fin. Scaly sheath reduced, present only on anterior

half of fin 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 it is reported by

these authors to occur commonly. However, many specimens supposedly of this

species may be C. melampygus. The area in which it 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 Islands0 5 cw.

Caranx) medusicola

BC60-488

specimen No. 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 fit 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 fin 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 fin in length 4.56; upper jaw in head, 2,52, Dorsal

VIII-I, 20; anal?II-I, 16; pectoral I, 19; ventrals I, 5. Gill rakers on lower limb of first 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. Gill rakers on lower limb of first arch 16-17. Depth in length 2.65-2.84. Head in length 2,67-3.31.

Pectoral fin 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. Size Range Cat. No. Specimens Locality (mm.)

BC58-398 1 East Coast of Florida 166 BC61-92 1 Florida Bay, Monroe, Florida 100 UMML408 1 Key Largo, Florida 178 UMML1150 1 300, 16• N; 80°, 21« W-Combat Stn. # 70 100 UMML1179 1 Dry Tortugas, Florida 146 UMML1457 1 Dry Tortugas, Florida 149 UMML2150 1 Long Key, Florida 184 UMML3048 1 Virginia Key, Florida 175 UMML7397 1 Virginia Key, Florida 156 UMML7784 1 Little Duck Key, Florida 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

first arch 26-28. Depth in length 2.53-3.49 (x 3.20). Head in length

3.51-3.75 (x 3.62). Pectoral fin 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 fin ray counts and colour pattern are extensively used in these keys. Overlap was found in all such characters for Caranx marginatus, melampygus and medusicola. The key given below separates all the species of Caranx known from the tropical east Pacific.

A Grill rakers on lower limb of first 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 fin Caranx caballus

AA Grill rakers on lower limb of first arch less than 22 (including rudiments)•

C Scales absent from breast except small diamond shaped patch in front of ventral fins. Caranx hippos

CC Breast scaled.

D Body all dark brown; front of head usually concave; anterior portion of dorsal and anal fin reaching well past l/2 fin base. Caranx lugubris

DD Body not all dark brown; front of head usually not concave; anterior portion of dorsal and anal fin not reaching well past 1/2 fin base.

E Head length over upper jaw length 2,00- 2.40 (usually between 2.05-2.25); body never spotted; gill rakers 17 or less, rarely 18; dorsal times anal fin rays less than 365; vertebrae 25. 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} gill rakers, 18 or more, rarely 17 or less; dorsal times anal fin rays greater than 365; vertebrae 24.

Anal fins pointed or almost so, longest anal ray 2\ or less in base of fin; anterio-dorsal profile not straight. Caranx melampygus

FP Anal fin round on anterior raised portion, longest anal ray greater than 2i in the fin 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, it 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 in the

Table below.

TABLE II

L L H Lateral Gill D H M Scutes Rakers

C. crysos 3.20 3.62 2.42 46.7 26.8 Average 2.53-3.49 3.51-3.75 2.24-2.53 45-51 26-28 range

C. caballus 3.78 3.77 2.68 44.1 28.6 Average 3.32-4.78 3.49-4.01 2.46-2,80 38-51 25-31 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 is 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) it is very likely that the C. crysos-caballus stock was - 40a -

I5n

crysos INDEX 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). head length fork length fork length 30-i upper jaw length head length depth

C caballus C crvsos 20-

£ 10- o> E o

C. marginatus

C. latus 20"

10-

0- i 2.0 2.5 3.0 "ST 4.0 2.5 3.0 35 4.0 4.5 Figure 21. - 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 it 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, gill raker count and scute count. C. vinctus does not fit Williams* (1958) Caranx generic restriction as it has two rows of teeth on the lower jaw and does not have falcate dorsal and anal fins. The origin of C, vinctus is difficult 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 all 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 gill raker count, more scutes on the lateral line and shorter ventral fins, but there is a great area of overlap in all 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 first 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 al, 1953) describes C, sexfasciatus from the Marshall and Mariannes Island as a different form making it 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. if it 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 Gill 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 all marginatus examined had 25 vertebrae and all latus

24 vertebrae and in all other Caranx species examined the vertebral count was constant within a given species, it 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 is distributed circumtropically around rocky islands.

Often, as in the Americas, it 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 it 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 Cleopha I., Tres Marias I" rS. 0 0 107-151 mm. 2 Socorro I., Revillagigedo I" 0 0 110-131 mm. 18 Clarion I., Revillagigedo Is* 0 0 119 mm. 1 Socorro I., Revillagigedo I 10-30 1 or 2 210-241 mm. 4 Socorro I., Revillagigedo Is] 50-100 100 396-670 mm. 5 Socorro I., Revillagigedo I 50-100 100 642-675 mm. 3 Maria Madre I.,Tres Marias I

From the examination of the specimens above (from 89-675 mm.) it 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 is absent from the Atlantic,

On this basis it 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 is 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 it 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 fin 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 melampygus 210mm.

/' C, medusicola 157mm

Figure 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 it is considered as such in this work. The origin of C. medusicola is difficult to ascertain. Morphologically it 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 is: 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 ft. 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> coral reef areas

Figure 23. EQUATORIAL PACIFIC OCEAN Barrier area indicated by label. - 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 little 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 first - 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: firstly, 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 if 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 all 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 little 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.

EAST PACIFIC INDO-PACIFIC BARRIER EAST PACIFIC ISTHMUS WEST ATLANTIC DIFFERENTIATION

C. sexfasciatus C. marginatus C. latus iC, marginatus from C. latus and C. sexfasciatus - 100$ on vertebral count

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 is, therefore, only 60$ effective when considering either east-west or west-east movement in the past. Today, however, it 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 it 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 is 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 is supported by most

of the literature and is not argued here; however, it 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 is 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 fin 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

it is unlikely that the postulated 3 to 4C° Pleistocene drop in temperature had a serious effect on the shore fish fauna. BIBLIOGRAPHY

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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 Appendix 2 - C. caballus data (All measurements in millimetres)

Head Pectoral Upper Eye Dorsal Anal Pect- Caudal Gill Length Depth Length Fin Jaw Length Fin Fin oral D M V Rakers Vertebrae Scutes m m Length Length - Fin

191.0 51.3 50. 0 61.4 19.4 12.7 VHI-I 24 II--118 -- 7 21 8 28 10+ 15 44 185. 5 49. 5 47.6 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 70.9 75.6 - 29. 5 - 22 19 - - 28? - 44 176. 1 46.9 46.2 50.8 16.7 12.0 22 18 - - 27 ? 10+ 15 45 284. 1 74.2 75.2 97.8 28.8 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 66.5 20. 1 - 24 19 8 20 7 30 10+ 15 46 196. 9 49.6 52.0 58.6 19.0 - 23 20 8 20 8 28 10+ 15 49 280. 1 67.8 71.2 92.0 26. 5 - 24 18 - - 29 - 39 264.7 69.0 69.9 85.8 26. 1 - 23 17 1-22 - - 28 - 41 167. 1 49. 5 44.4 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 75.7 73. 5 98.7 27.9 - 22? 20 1-23 - - 26? - 43 204.7 57.2 54.0 - 19.6 - 23 19 - - 28 - 42 211.8 57.2 54.7 63.2 20.8 - 23 19 1-22 - - 28 - 42 226.6 64. 1 58.0 67.4 21.6 - 24 20 - - 30 - 43 241.2 65.9 60. 1 72.4 22. 5 - 22 19 1-22 - - 29 - 46 272. 5 76.0 72. 5 92.8 27.8 - 23 19 - 28 - - 326. 5 86.7 85.0 108.8 32. 1 - 22 18 - - 29, - 46 350.9 85.0 94.5 127.4 36.0 - 24 20 - - 30 - - 199.6 45.7 51.3 60.3 19.4 12.0 21 18 8 20 7 29 10+ 15 44 310.6 64.9 78. 3 87.0 29.2 - 24 19 - - 29 - 43 293.7 77.0 82.0 99.5 30.4 - 23 18 - - 29 - 46 168.9 46.7 45.9 50.9 17.4 - 23 ? 16? - - - 25 - 45 283. 8 69.6 75.0 87.7 27.9 16.4 24 20 - - 30 - 44 262. 0 67.0 •67.6 85.7 25.9 17.0 22 21 - - 29 10+ 15 45 195.4 48.8 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 Gill Length Fin Jaw Length Fin Fin oral D V Rakers Vertebrae Scute M m Length Length Fin m

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^ers 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. U5: 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

V Length Depth Length Fin Jaw Length Fin Fin oral m M 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 10.2 23 17 - - 17 _ 33 142.9 49.6 43. 1 45.0 19.4 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 16 16 10+ 14 - 20 17 1-22 8 18 8 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 Length Depth Length Finn Jaw Length Fin Fin oral D^ M V^ Rakers Vertebrae Scutes Length Length Fin

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 Gill Length Depth Length ,Pin,, Jaw Length Fin Fin oral D M V Bakers Verte- Scutes en® 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 Gill Length Depth Length Fin Jaw Length Fin Fin oral D M V Rakers Verte- Scutes Length Length Fin brae

150.0 55.9 49.4 50.8 21.6 12.8 VIII I 20 11-116 22 8 18 8 16 10+15 33 173.1 61.2 52.6 — 23.7 14.7 20 16 22 7 19 9 16 10+15 33 153.0 54.1 45.3 48.0 21.3 13.1 20 17 20 — 18 10+15 34 151.9 23.7 47.1 48.5 20.6 12.8 20 16 22 — 16 10+15 31 149.9 53.6 44.1 46.8 20.0 12.4 20 15 21 — 17 10+15 32

Appendix 7 - C. latus data ]

160.0 60.0 48.4 48.0 23.9 13.0 VIII I 20 II-I 18 20 8 17 7 17 10+14 36 112.0 39.5 34.1 29.3 14.8 9.4 20 17 19 9 16 8 17 10+14 38 87.9 32.0 26.8 24.8 12.9 8.4 21 16 20 8 17 8 16 10+14 35 53.2 20.1 19.9 12.6 7.5 5.4 20 17 — —i 16 10+14 33 85.0 30.9 26.9 22.2 10.9 7.0 21 16 21 3 \% 7 16 10+14 33

\ Appendix 8 - C. lugubris data

Head Pectoral Upper Eye Dorsal Anal Pect• Caudal Gill Length Pin Length Pin Pin oral D M V Rakers Verte• Scutes Length Depth Jaw ra ra Length Length Fin brae

_ 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 Gill Verte• Length Pin Jaw Length - Pin Pin oral D M V Rakers brae Scutes Length Depth m Length Length Pin iimi

129.2 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 ^ 111.4 43.8 31.9 35.0 12.6 9.6 VIII-I 22 20 — — 19 10+14 32 ? 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 — 22 21 9 32 114.8 42.7 31.3 35.0 12.8 8.9 19 8 17 } 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 Gill 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