A Systematic Monograph of the

Tongue Soles of the Genus

Cynoglossus Hamilton-Buchanan

(Pisces: Cynoglossidae)

A. G. K..MENON

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 238
SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of "diffusing knowledge" was expressed

by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph

Henry outlined a program that included the following statement: "It is proposed to

publish a series of reports, giving an account of the new discoveries in science, and

of the changes made from year to year in all branches of knowledge." This theme

of basic research has been adhered to through the years by thousands of titles issued

in series publications under the Smithsonian imprint, commencing with Smithsonian

Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoology

Smithsonian Studies in Air and Space

Smithsonian Studies in History and Technology

In these series, the Institution publishes small papers and full-scale monographs

that report the research and collections of its various museums and bureaux or of

professional colleagues in the world cf science and scholarship. The publications are

distributed by mailing lists to libraries, universities, and similar institutions throughout

the world.

Papers or monographs submitted for series publication are received by the

Smithsonian Institution Press, subject to its own review for format and style, only

through departments of the various Smithsonian museums or bureaux, where the

manuscripts are given substantive review. Press requirements for manuscript and art

preparation are outlined on the inside back cover.

S. Dillon Ripley

Secretary

Smithsonian Institution
SMITHSONIA N CONTRIBUTION S TO ZOOLOGY • NUMBE R 238

A Systematic Monograph of the

Tongue Soles of the Genus

Cynoglossus Hamilton-Buchanan

(Pisces: Cynoglossidae)

A. G. K. Menon

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRAC T

Menon, A. G. K. A Systematic Monograph of the Tongue Soles of the Genus

Cynoglossus Hamilton-Buchanan (Pisces: Cynoglossidae). Smithsonian Contributions to Zoology, number 238, 129 pages, 48 figures, 21 plates, 1977.—The

genus Cynoglossus Hamilton-Buchanan is characterized, and its relationship with

other cynoglossid genera is discussed. Forty-nine species including a new species,

C. lachneri, are recognized.

The origin, distribution, and evolution of the genus are discussed. The genus

is of tropical Indo-Pacific and eastern tropical Atlantic in distribution, and the

members are mostly marine sandy or muddy shallow-water inhabitants. From

the present-day distributional pattern, the genus is considered to have evolved

in the Indo-Malayan Archipelago during the Pliocene, and the geological

evidences providing explanation for its dispersal to the eastern tropical Atlantic,

presumably, during the upper Pliocene have been reviewed.

The genus is composed of an extremely homogeneous assemblage of highly

specialized species. Based on a comparative study of the epicranial bony system

in different families of flatfishes, it is concluded that in the genus Cynoglossus,

it is modified to suit a burrowing habit.

The apparently primitive and specialized characters in the genus are also

outlined. The possession of a lateral line on the blind side is considered a

primitive character, for it serves no purpose to a fish adapted for a burrowing

mode of life. Thus, the absence of a lateral line on the blind side, reduction in

the number of caudal fin rays, reduced size of scales, contiguous or closely set

eyes with narrow interorbital space, or reduced or minute eyes with wide interorbital space, and loss of posterior slitlike nasal opening are considered specialized characters necessary for a burrowing habit. Based on the morphological

features and on zoogeography, the 49 species have been divided into six groups

and 17 complexes, and their hypothetical evolutionary relationships are traced.

A key for the separation of species is provided; separate keys for the groups

and complexes are not provided, however, since they were mainly based on the

pattern of distribution and hence arbitrary in nature. Under each species, the

complete synonymy, description, coloration in preservation, size diagnosis and

affinities, note on synonymy, information on the type and other material

examined, and geographical distribution are given. Lectotypes (wherever necessary) have been selected. Lectotype selection has also been made (wherever

necessary), even to a nominal species now considered as a junior synonym to

avoid any confusion in the future as to its true identity. The species are illustrated in 40 outline drawings and 21 plates. Eight distributional maps are also

provided.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Menon, A. G. K.

A systematic monograph of the tongue soles of the genus Cynoglossus Hamilton-Buchanan

(Pisces, Cynoglossidae)

(Smithsonian contributions to zoology ; no. 238)

Bibliography: p.

1. Cynoglossus. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 238.

QL1JS54 no. 238 [QL638.C93] 591'.08s [597.5] 76-608109
Contents

Page

Introduction 1

Acknowledgments 1

Methods 3

Definition of Measurements and Counts 3

Radiography 3

Clearing and Staining 3

Family CYNOGLOSSIDAE 3

The Relationships of the Genera 6

Key to the Genera of Cynoglossidae 6

The Lateral-Line System 6

The Scales 10

The Epicranial Bony System 11

The Species Groups and Complexes and Their Hypothetical

Evolutionary Relationships 13

Genus Cynoglossus Hamilton-Buchanan, 1822 16

Zoogeography and Evolution 18

Distribution 18

Geological History of the Red Sea 25

Dispersal during the Pliocene to Recent 25

Key to the Species 26

The canariensis group 28

The canariensis complex 28

1. C. canariensis Steindachner 28

2. C. monodi Chabanaud 30

3. C. senegalensis (Kaup) 31

4. C. dubius Day 32

5. C. borneensis (Bleeker) 34

The browni complex 35

6. C. browni Chabanaud 35

The bilineatus complex 36

7. C. bilineatus (Lace'pede) 36

The attenuatus complex 38

8. C. attenuatus Gilchrist 39

9. C. lachneri, new species 40

10. C. dispar Day 40

The kopsi group 42

The kopsi complex 42

11. C. kopsi (Bleeker) 42

12. C. interruptus Gunther 45

13. C. joyneri Gtinther 46

The itinus complex 48

14. C. itinus (Snyder) 48

The ogilbyi complex 49

iii
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page

15. C. ogilbyi Norman 49

16. C. maculipinnis Rendahl 50

17. C. broadhursti Waite 51

The ecaudatus complex 51

18. C. ecaudatus Gilchrist 52

19. C. cadenati Chabanaud 53

20. C. dollfusi (Chabanaud) 54

21. C. sinusarabici (Chabanaud) 55

The sealarki complex 56

22. C. sealarki Regan 56

23. C. microphthalmus (Bonde) 57

24. C. zanzibarensis Norman 58

25. C. capensis (Kaup) 59

The arel group 59

The arel complex 60

26. C. arel (Schneider) 60

27. C. robustus Giinther 64

28. C. lingua Hamilton-Buchanan 65

The cynoglossus group 67

The cynoglossus complex 67

29. C. cynoglossus (Hamilton-Buchanan) 68

30. C. semifasciatus Day 70

31. C. macrostomus Norman 72

The monopus complex 73

32. C. monopus (Bleeker) 73

The puncticeps complex 75

33. C. puncticeps (Richardson) 75

34. C. durbanensis Regan 79

35. C. gilchristi Regan 80

The lida complex 81

36. C. lida (Bleeker) 81

The carpenteri group 82

The carpenteri complex 83

37. C carpenteri Alcock 83

38. C. acutirostrs Norman 84

39. C. marleyi Regan 85

40. C. suyeni Fowler 86

The heterolepis group 87

The heterolepis complex 87

41. C. heterolepis Weber 87

42. C. feldmanni (Bleeker) 89

43. C. waandersi (Bleeker) 90

44. C. kapuasensis Fowler 91

45. C. semilaevis Giinther 92

46. C. abbreviatus (Gray) 93

47. C gracilis Giinther 95

48. C. microlepis (Bleeker) 97

The macrophthalmus complex 99

49. C. macrophthalmus Norman 99

Literature Cited 100

Plates 109
A Systematic Monograph of the

Tongue Soles of the Genus

Cynoglossus Hamilton-Buchanan

(Pisces: Cynoglossidae)

A. G. K. Menon

Introduction

The present work was begun during 1967-1968

at the Division of Fishes of the National Museum of

Natural History, Smithsonian Institution, Washington, D.C., where I was employed for a year as a

Senior Postdoctoral Visiting Research Associate and

continued for short periods during OctoberNovember 1968 at the British Museum, London,

and Museum National d'Histoire Naturelle, Paris,

and later at the Zoological Survey of India, Calcutta. The opportunities thus afforded for the examination of material were exceptional, and a much

larger number of specimens have passed through

my hands than I had at first thought possible. The

results of my studies are presented here and I hope,

if opportunity is given, to pursue in more detail

behavioral, ecological, and life history studies of

the cynoglossid flatfishes in a subsequent work.

The genus Cynoglossus Hamilton-Buchanan is

tropical Indo-Pacific and eastern tropical Atlantic

in distribution, and its members occur along sandy

or muddy shores and estuaries. The species are

marine or estuarine shallow-water burrowing forms

with a few species (C. microlepis (Bleeker), C. heterolepis Weber, and C. kapuasensis Fowler) enterA. G. K. Menon, Zoological Survey of India, Southern Regional Station, 69 Santhome High Road, Madras 600028,

India.

ing fresh-water streams. Of the 49 species included

in the genus, one, Cynoglossus lachneri Menon, is

described here as new.

No comprehensive study of the genus Cynoglossus

has so far been made, although Norman (1928),

Weber and de Beaufort (1929), Punpoka (1964), and

Shen (1967, 1969) revised certain Indo-Pacific species and Chabanaud (1949a-d) and Cadenat (1960)

revised certain species of the Atlantic. The present

work is based on specimens available to earlier

workers and on a large number of additional specimens resulting from intensive collecting in recent

years, especially in the Indian Ocean during the

International Indian Ocean Expedition (1960-

1965). It is the first attempt to study the systematics

of the genus in the strict sense understood by

Myers (1952).

ACKNOWLEDGMENTS.—A large number of individuals and institutions aided me in many ways, especially by the loan of specimens or by making

them available for study, by provision of laboratory

space or by generously giving information about

specimens, notes on specimens in their care, or

data from writings unavailable to me, or by helping freely in the translation into English of some

of the papers written in French. To each of them

I am gratefully indebted, for without their consideration my study could never have been accomplished. To the institutions and their present or

former personnel, responsible for the assistance I

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

received, I extend my sincere thanks. A list of those

individuals and institutions follows, together with

the appropriate abbreviation used throughout the

text.

AMNH American Museum of Natural History, New York,

D. E. Rosen

AMS Australian Museum, Sydney, F. H. Talbot, J. R.

Paxton

ANSP Academy of Natural Sciences of Philadelphia, Philadelphia, J. E. Bolhke, J. C. Tyler

BMNH British Museum (Natural History), London, E.

Trewavas, P. H. Greenwood, N. B. Marshall,

P. J. P. Whitehead, A. C. Wheeler

CAS California Academy of Sciences, San Francisco,

W. N. Eschmeyer (George Vanderbilt Foundation

collection has now been transferred to the Californian Academy of Sciences and bears the abbreviation "GVF" in parentheses after the

abbreviation "CAS")

CU Cornell University, Ithaca, New York, E. C. Raney

HUJ Hebrew University, Jerusalem, the late H. Steinitz

KU Kyoto University, Japan

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, M. M. Dick

MNHP Museum National d'Histoire Naturelle, Paris, M. L.

Bauchot

NHV Naturhistorisches Museum, Vienna, P. Kahsbauer

NMNH National Museum of Natural History, Smithsonian

Institution, Washington, D.C.

RMNH Rijksmuseum van Natuurlijke Historie, Leiden, M.

Boeseman

RU Rhodes University, Department of Ichthyology,

Grahamstown, South Africa, the late J. L. B.

Smith, Mrs. M. M. Smith

SOSC Smithsonian Oceanographic Sorting Center, Washington, D.C, L. W. Knapp

UMMZ University of Michigan, Museum of Zoology, Ann

Arbor, Michigan, R. R. Miller, R. M. Bailey

USNM Acronym for former United States National Museum

collections in NMNH

UZMK Universitetets Zoologiske Museum, Copenhagen, J.

Nielsen

WAM Western Australian Museum, Perth, R. McKay

ZMA Zoologisch Museum, Amsterdam, N. Nijssen

ZMB Zoologische Museum, Berlin, C. Karrer

ZSI Zoological Survey of India, Calcutta

To the following from the National Museum of

Natural History, Smithsonian Institution, Washington, D.C. I am deeply indebted: Dr. Ernest A.

Lachner, Curator of Fishes, Division of Fishes, for

initiating this project and for constant guidance

and encouragement; Drs. Daniel M. Cohen and

Bruce B. Collette of the National Marine Fisheries

Service Systematics Laboratory for their considerable

help and constant encouragement throughout the

period of my work in the Division of Fishes. I am

most grateful to Drs. E. Trewavas and P. H.

Greenwood and through them the Trustees of the

British Museum (Natural History) for providing

me with the outline drawings of the various species of Cynoglossus prepared by Mr. Hubert Williams under the supervision of the late Dr. P.

Chabanaud and for permitting me to use them

in the present work. To Mr. P. J. P. Whitehead

I am particularly grateful for his kindness in

examining on my behalf some of the types in the

British Museum, for attending to my several queries

in the course of the preparation of this work, and

for carefully and critically going through the manuscript and offering valuable suggestions for the

improvement of the work. Dr. Boeseman, Curator

of Fishes, Rijks Museum van Natuurlijke Historie, Leiden, searched the literature, as well as the

collections in Leiden, and helped me to select

the appropriate types for species of Cynoglossus

described by Bleeker. But for the very kind and

generous help that he readily extended to me this

work could not have been completed, and it is very

gratefully acknowledged here. During the course

of my work I have had the benefit of personal discussions with my colleagues in the National Museum of Natural History, especially Drs. R. H.

Gibbs, W. R. Taylor, V. Springer, S. Springer, S. H.

Weitzman, L. Schultz, W. Aron, L. W. Knapp,

and R. Kanazawa. To all of them I am grateful.

I have also benefitted from personal discussions with

Dr. Carl L. Hubbs on the adaptation of Cynoglossus

for a burrowing habit.

The staff of the Central Photo Laboratory of the

National Museum of Natural History made the

photographs. In the X-ray room of the Division

of Fishes (NMNH), the radiographs were made by

me, with the help of Mr. E. N. Gramblin.

I should like to record my gratitude to the National Research Council (National Academy of

Sciences and Engineering) of the United States,

Washington, D.C, for awarding me a Smithsonian

Senior Postdoctoral Visiting Research Associateship

to enable me to work for a year in the Division of

Fishes (NMNH). To the Director of the Zoological

Survey of India and the Ministry of Education and

Youth Services, Government of India, I am thankful for the grant of study leave to enable my visit

to the United States.

My wife, Mrs. Rema G. Menon, who came with

me to the United States, helped me a great deal by
NUMBER 238

regularly accompanying me to the laboratory and

making counts and measurements.

Methods

DEFINITION OF MEASUREMENTS AND COUNTS.—

Measurements were taken on the ocular side of

each specimen, using a pair of needlepoint dividers

and recording to the nearest one-tenth of a millimeter.

Standard Length: Taken from the tip of the

snout to the midlateral posterior margin of the

hypurals as indicated externally in the specimen.

Head Length: Taken from the tip of the snout

to the posteriormost point reached by the fleshy

margin of the opercle.

Diameter of Eye: The vertical diameter taken

between the dorsal and ventral rims of the fixed

eye.

Snout Length: Taken from the tip of the snout

to the anterior rim of the fixed eye.

Interorbital Width: Taken as the nearest distance between the upper rim of each orbit whereever the eyes are separate.

Distance between Snout and Corner of Mouth:

Taken in a direct line, being the distance between

the tip of the snout and the posterior corner of the

cleft of the mouth (measured by keeping the needle

of one arm of the divider inside the mouth cleft

and the other at the end of the snout).

Distance between Corner of Mouth and Gill

Opening: Taken in a direct line, being the distance

between the inside corner of the cleft of the mouth

and the posteriormost point reached by the fleshy

margin of the opercle.

The enumeration of dorsal and anal fin rays in

Cynoglossus is most difficult and tedious, and much

of the confusion in the taxonomy of Cynoglossus

has been cauesd by erroneous counts made on a

limited number of specimens of a particular species. The counts of the dorsal and anal fin rays in

this work were taken, therefore, from radiographs

and some were from cleared and stained specimens.

Caudal Rays: All rays in the caudal fin. Since the

caudal fin is confluent with both the dorsal and the

anal fin some difficulty was experienced in the

enumeration of the caudal rays. The caudal rays

were therefore counted by placing the fin on the

glass stage of a binocular microscope and projecting strong light through the fin. Removing the

mucus and skin by means of a needle, a count was

taken and then the fin was turned over and the

enumeration checked on the blind side.

Vertebrae: In Cynoglossus the last six abdominal

vertebrae bear short haemapophyses (Figure 1). The

caudal vertebrae are, therefore, those beginning

from the first interhaemal spine, which forms the

posterior boundary of the abdominal cavity. In

radiographs the position of the first interhaemal

spine can be readily seen.

RADIOGRAPHY.—Radiographs were prepared by

the use of soft X-rays. In this method the fishes

were taped closely to the film holder containing

high contrast film and exposed at 22 kV. 125

milliamp-seconds, with the X-ray unit focussed

about 20 inches from the object.

CLEARING AND STAINING.—A few specimens were

cleared and stained for osteological studies and for

checking counts made from radiographs. Different

methods of clearing and staining specimens have

been used (Taylor, 1967). This above method is

successful when used on fresh or newly preserved

specimens but not on specimens stored in various

preservatives for several years (which is most often

the case in museum collections). Such specimens

may remain opaque because of dark stains in the

flesh or they may swell, rupturing the membranes

during the clearing process. In order to overcome

these difficulties Taylor (1967) evolved a new

method, substituting enzyme digestion for alkaline

maceration of tissues.

In the improved enzyme method of Taylor, trypsin powder (Fisher Scientific Company's "Purified

Trypsin Powder," 1 : 80 MF) is used in sodium

borate (borax) buffer solution, which has proved

very effective in so far as it supports the most rapid

enzyme activity while maintaining a relatively stable and desirable pH over a long period of time.

This method has been successfully used in the

present work.

Family CYNOGLOSSIDAE

The Heterosomata to which the Cynoglossidae

belong can be divided into seven families: Psettodidae, Citharidae, Scopthalmidae, Bothidae, Pleuronectidae, Soleidae (true soles), and Cynoglossidae

(tongue soles or solelike fishes with eyes on the left

side). The soles (Soleidae and Cynoglosside) are

distinguished by having a small mouth, the lower
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

jaw not prominent, the jaws of the blind side

strongly curved and toothed, no preopercular margin, the gill membranes fused with free branchiostegal rays, the symmetrical position of the nasal

organs, the absence of a postcleithrum in the pectoral arch, and the absence of ribs. In spite of these

common characteristics Norman (1934:38) doubted

whether the two families are really closely related.

Norman cited the example of certain Australian

and New-Zealandian genera of the Pleuronectid

subfamily Rhombosoleinae (Ammotretis, Colistium,

Peltorhamphus) in which some of the species exhibit a strong general resemblance to members of

the Soleidae. Especially in Colistium the general

form of the body, the shape of the head, particularly its preorbital part, the small eyes, the symmetrical nasal organs, the strongly curved jaws of

the blind side, the absence of teeth in those of the

ocular side, and the extension of the dorsal fin to

the end of the snout are all characteristics found in

the members of the Soleidae. The development of

membranous folds on the blind side of the rays

of the marginal fins, the modification of many of

the scales on the blind side of the head to form

filamentous processes, and the fringed lower lip are

some of the other soleid features developed by the

Rhombosoleinae (Norman, 1926:259). Chabanaud

(1933, 1934b, 1936, 1937) gave much consideration

to the probable polyphylelic origin of the Soleoidea from other flatfishes. Kyle (1921:119-121) believed that the flounders and the soles, and even

the divisions within each of these groups, have had

separate origins from symmetrical fishes.

Although it seems probable that the origin of

the unique asymmetry of flatfishes was a single

evolutionary event, it must be admitted that the

Heterosomata as a whole are held together by

little more than the single character that the two

eyes are on one side of the head. Many of the

features in which the several families agree now

appear to represent convergent adaptations (Hubbs,

1945). The work of Parker (1903) on optic nerves

of the Heterosomata is, however, of special significance in this connection. In ordinary bony fishes the

optic chiasma is dimorphic in character, the right

nerve crossing over the left about as often as the

left over the right. In the families Soleidae, Cynoglossidae, and Psettodidae, the chiasma is dimorphic, with the nerve of either the left or the right

eye the more dorsal in the optic chiasma. In all

other flatfishes, whether dextral or sinistral, the

nerve of the migrating eye is dorsal. In the genera

of flounders, which are normally dextral (with eyes

and color on the right side), the left nerve crosses

over the right in all individuals, even in reversed

specimens. Similarly in the normally sinistral groups

the right nerve is superior even though the individual is variant in having the eyes on the right side.

As a result the chiasma is characterized as partly

uncrossed in normal individual but doubly crossed

in the reversed specimens.

Parker (1903) interpreted the correlation of the

type of chiasma with the position of the eye in

flounders as adaptive, for when the nerve of the

migrating eye is dorsal the chiasma is partly uncrossed rather than doubly crossed as it is when

the nerve of the migrating eye is ventral in the

chiasma. The chiasma remains dimorphic, however,

in both dextral and sinistral soles. This can be

explained as due to the relative development of

the optic nerves in the two groups. In the flounders, for example, the optic lobes and nerves are

larger than in the soles (Evans, 1937:309-310) and

are much more conspicuous than the olfactory

nerves, occupying a large part of the cavity in

which they lie. A complicated arrangement (double crossing) of the nerves may, therefore, involve

a mechanical or developmental disadvantage. In

soles, on the other hand, the optic nerves are tiny

strands lying loose in an extensive space under the

large olfactory nerves. In these fishes the olfactory

and tactile senses are presumably better developed

than the visual. Thus, the double twist of the optic

nerves in half the individuals of each species of

sole has apparently not been of sufficient selectional

significance in the soles, as it has in the flounders,

to a fixation of the optic chiasma type in correlation with the usual position of the eye.

The soles, in having several characters in common including the primitive dimorphic type of

chiasma, form a natural group, probably split off

very early from the other Heterosomata.

The members of the family Cynoglossidae are

typically sinistral, with a dimorphic optic chiasma;

the jaws are strongly asymmetrical; usually there

are two nares on each side of the head, the anterior

one tubular, the narial tube of the eyed side always arising in front of the fixed eye; the dorsal

and anal fins are confluent with the caudal; the

dorsal fin extends onto the head parallel to the
NUMBER 238

IN

NS DR

E-f—-;

RCFIGURE 1.—Skeleton of C. puncticeps, drawing of a cleared and stained preparation from a

109.0 mm SL specimen (USNM 109799) from the Gulf of Thailand: a, entire skeleton; b, neurocranium; c, epicranium; d, last three abdominal and first two caudal vertebrae; e, epural and

hypural bones supporting the ten caudal rays. (AIS = anterior pseudointerneural spine, C =

cranium, E = erisma. In = in tei neural spine, PIS = posterior pseudointerneural spine,

RC = rostral cartilage, AR = anal fin rays, CR = caudal fin rays, DR = dorsal fin rays,

EP = epural bone, HP = hypural bone, HS = haemal spine, IH = interhaemal spine,

NS = neural spine.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

axis of the cranium, the first dorsal ray never being

inserted behind the vertical from the posterior

margin of the eye, this ray remaining above the

level of the migratory eye even when the dorsal

fin is extended to the tip of the snout; the pectoral

fins are lacking in the adult; the pelvic of the eyed

side is also lacking (except in rare individual cases

where it is situated on the eyed side much above

the midventral line, as in the case of Cynoglossus

Zanzibarensis, Figure 25); the pelvic fin of the blind

side is present with four rays always inserted on

the midventral line; the anus and the opening of

the oviduct are on the blind side; the urinary

papilla is long and situated midventrally in front

of and attached to the first anal fin ray; the scales

are generally ctenoid; the tactile fringes of the

lower side of the head are either short or absent

and are replaced by epidermal thickness; and there

are no epidermal hairs.

The rostral process of the neurocranium is absent or rudimentary; abdominal vertebrae are

usually 9, sometimes 10, or even rarely 12 (C. robustus, C. abbreviatus, C. semilaevis); the number

of caudal vertebrae is 33 to 66; the neural arch

is complete in all the vertebrae, including the first

abdominal vertebra; the haemal arches are complete

as far forward as the fourth abdominal vertebra;

and an anal interhaemal spine is present, attached

to the first caudal haemal spine near its extremity.

The digestive and urinary organs do not extend

to the caudal region, only the ovaries and the

posturethral portion of the urinary bladder occupy

the caudal region, and a swim bladder is absent in

the adult.

The family Cynoglossidae though comprising

over 100 species is a very homogeneous group,

which is evident from the fact that it is comprised

of only three genera: Symphurus Rafinesque 1810;

Cynoglossus Hamilton-Buchanan, 1822; Paraplagusia Bleeker, 1886. The genera Symphurus and Cynoglossus contain most of the species; Paraplagusia

consists of only four species, P. bilineatus (Bloch),

P. blochi (Bleeker), P. japonica (Temminck and

Schlegel), and P. guttata (Macleay). The three genera are so homogeneous that their subdivision into

subgenera would appear only artificial. In spite of

its evident homogeneity, the family Cynoglossidae

is divisible into two subfamilies, the Symphurinae

consisting of the genus Symphurus and the Cynoglossinae consisting of the genera Cynoglossus and

Paraplagusia. The two subfamilies are perfectly

distinguishable from the point of view of their respective morphology and the geographical distribution.

THE RELATIONSHIPS OF THE GENERA

During the course of this revision an examination was also made of the four species of Paraplagusia, as well as a dozen species of Symphurus.

Skeletonized preparations of C. puncticeps, C.

browni, C. heterolepis, C. robustus, C. lida, Paraplagusia blochi, and a species of Symphurus were

studied.

Paraplagusia can be distinguished from Cynoglossus mainly by its possession of a series of fringes

on the lips on the ocular side. In all other features,

including the osteology, Paraplagusia is very similar to Cynoglossus. In the well-developed and more

strongly bent erisma and strongly hooked snout,

the tip reaching the rear of the lower eye or even

beyond, Paraplagusia is considered as a form more

highly specialized than Cynoglossus for a burrowing

habit.

Key to the Genera of Cynoglossidae

1. Ventral fins connected with anal, lateral line on ocular side, snout hooked, mouth inferior 2

Ventral free from anal, no lateral line on ocular side, snout not hooked, mouth anterior Symphurus

2. Lips with fringes, 2 or 3 lateral lines on ocular side Paraplagusia

lips without fringes, 1, 2, or 3 lateral lines on ocular side Cynoglossus

THE LATERAL-LINE SYSTEM

Except for Symphurus, the other members of

the family Cynoglossidae have the lateral-line system well developed.

Head Region: The head region of all the species

of the Cynoglossinae has a complex system of canals. In all the species a supraorbital canal is

present, which extends from the snout through the
NUMBER 238

TABLE 1.—Comparison of subfamilies Symphurinae and Cynoglossinae

Characters

Snout

Mouth

Jaws and teeth

Scales

Lateral line

Pelvic fin

Pectoral fin

Urinary papilla

Neurocranium

Epicranium

Vertebrae

Symphurinae

Not hooked

Terminal

On ocular side premaxillary

smooth, but dentary armed

with series of needle-shaped

teeth; jaws on blind side

toothed, premaxillary and

dentary teeth arranged in small

number of longitudinal series,

forming long narrow band

Ctenoid; head scales of blind

side imbricate

Absent on both sides

Free from anal fin

Rudimentary

Short, attached to first anal ray

Vault of cranium absent due to

huge fontanelle; otic capsules

bulky and hemispherical; a

spinuous divergence backwardly

directed and attached to base

of anterior pseudointerneural

spine; supraoccipital cartilagenous (Chabanaud, 1940)

Anterior pseudointerneural spine

slender; erisma short, archlike

in shape; rostral cartilage not

developed

(9-10) + (38-42) = 57-52

All vertebrae bear diapophyses

that are heavy in abdominal

region and anterior half of

caudal region, but become

weaker posteriorly; abdominal

haemal spines begin with

fourth abdominal vertebra, and

are shorter and more strongly

oblique than those of caudal

series; abdominal haemal spines

gradually decrease in length

Cynoglossinae

Hooked

Inferior

Jaws on blind side only armed

with needle-shaped teeth; on

dentary and premaxilla they

form wide short band

Generally ctenoid; head scales

on blind side embedded;

scales in midlateral line

imbricate but with deep

median cleft and usually in

species with ctenoid scales

lateral line scales are provided with cteni on either

side of pore of lateral line

Well developed on ocular side;

midlateral line that, except

in C. sinusarabici, is

accompanied by marginodorsal line and frequently by

marginoventral line

Confluent with anal fin

Absent

Long, attached to first anal ray

Vault of cranium complete; otic

capsules not prominent;

prefrontal bone without

spinous divergence; supraoccipital calcified

(Chabanaud, 1940)

Anterior pseudointerneural

spine robust, quadrangular

or swordlike; erisma elongate,

falciform directly supporting

numerous interneural spines

advanced into cranium; rostral

cartilage well developed

(9-10 or 11, rarely 12) +

(33-66) = 42-76 or 77 or 78

Vertebral diapophyses absent

or present only the anterior

part of caudal region;

abdominal haemal spines begin with fourth abdominal

vertebra and are shorter and

more strongly oblique than

those of caudal series; all

abdominal and caudal vertebrae have dorsal spines
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Comparison of subfamilies Symphurinae and Cynoglossinae—Continued

Characters

Cleithrum

Brain

Larval development

Ecology

Symphurinae

from ninth to fourth and are

also gradually inclined more

and more backward; all

abdominal and caudal vertebrae

have dorsal spines; they are

thicker and stronger and less

inclined backward toward anterior end; neural spine of first

vertebra leans forward so much

that it is in contact with

posterior face of skull, that of

second vertebra distinctly

inclined forward but third one

only slightly

Smooth and weakly arched

Diencephalon not enlarged

(Ochiai, 1963:100)

Symmetrical larval stage known

(Kyle, 1913, 1921), with swimbladder on ocular hemisome

Marine but a single species (S.

orientalis (Bleeker) is reported

from freshwater at Peiping);

majority of species * are

eurybathyal (Goode and Bean,

1885. 1886; Alcock, 1889; GilChrist, 1889; Weber and de Beaufort, 1929, Ginsberg, 1951) and

range from 300 m to a

maximum 1964 m

Cynoglossinae

Subangularly arched; at its base

cuticular crest of coracoid

segment is hypertrophied into

an acute-angled apophysis

Diencephalon enlarged

(Ochiai, 1963:68)

Symmetrical larval stage known

(Seshappa and Bhimachar,

1955), with swimbladder on

ocular hemisome

Marine but a few species (C.

microlepis, C. heterolepis,

C. kapuasensis, C. feldmanni,

C. waandersi) are known to

occur in fresh waters; shallow

water burrowing forms

*Symphurus pusillus (Goode and Bean), 311 m; 5. septemstriatus (Alcock), 402 m; S. nebulosus

(Goode and Bean), 414 m; S. nigrescens Rafinesque, 420 m; S. niger (Goode and Bean), 457 m;

S. marginatus (Goode and Bean), 587 m; S. ftiscus Brauer, 638 m; S. macrophthalmus Norman,

640 m; S. ocellatus Bonde, 640 m; 5. striatus Gilbert, 730 m; S. trifasciatus Alcock, 782 m; S.

regani Weber and de Beaufort, 794 m; S. variegatus (Gilchrist), 823 m; S. woodsmasoni (Alcock), 896 m; S. gilesi (Alcock), 1080 m; S. australis Maculloch, 1464 m.

area above the eye and is connected to the midlateral line in the posterior part of the head. Above

the supraorbital, the cephalodorsal canal commences at the snout, runs posteriorward along the

dorsal edge of the head, and is connected to the

supraorbital commissure (a dorsal branch of the

supraorbital line at the posterior end of the head).

These two canals are invariably developed in all

the species of Cynoglossinae. From the posterior

end of the supraorbital canal the preopercular

canal commences, running ventrally toward the preopercular region. In some forms it turns and runs

forward to the corner of the mouth, while in others

it turns backward and runs to the edge of the

opercular region. From the anterior end of the

lower jaw to the operculum stretches the mandibuloopercular canal. At its posterior end it turns

upward and is connected to the preopercular canal
NUMBER 238

in some (C. bilineatus), whereas in other groups

the mandibuloopercular canal gives off a dorsal

branch at the posterior corner of the mouth, thereby

connecting it to the preopercular canal (C. abbreviatus). In certain other species the mandibuloopercular canal is not connected with the preopercular canal (C. interruptus). On the ocular

side there is also a small preorbital canal that starts

from the tip of the snout and extends obliquely

upward in a straight line to before or below the

upper eye.

On the blind side of the head the lateral-line

system is poorly developed or totally absent. Only

small traces of the supraorbital, cephalodorsalic,

or preopercular canals are found in some forms like

the bilineatus and the canariensis complexes,

where the lateral-line system of the abdominocaudal

region of the blind side is also developed.1

Abdominocaudal Region: The Cynoglossinae are

the only flatfishes that possess many lateral lines

(with the exception of C. sinusarabici, in which

there is only a midlateral line). All the other members of both Cynoglossus and Paraplagusia have

two or three lateral lines on the ocular side. On

the blind side they are generally fewer in number

than on the eyed side or are totally absent. In C.

dubius Day, C. monodi Chabanaud, C. canariensis

Steindachner, C. senegalensis Kaup, and C. borneensis (Bleeker) there is only one lateral line on the

blind side, while in C. bilineatus (Lacepede), C.

attenuatus Gilchrist, C. lachneri Menon, and C.

dispar Day there are two lateral lines on the blind

side. In all other members of Cynoglossus the

lateral-line system is totally absent on the blind

side.

Of the lateral lines on the ocular side the most

'In the Soleidae the lateral-line system of the blind side

of the head region is well developed and various modifications of the scales of the lateral line are found. In Aseraggodes a row of thin cutaneous flaps is developed on the

lateral line, the edges of which are serrated. In Heteromycteris and Soleichthys the cutaneous structures are hairshaped and in the latter they are arranged on the lateral

line like a brush. In Synaptura a well-serrated cutaneous

flap occurs, in which the lateral-line canals are embedded

(Ochiai, 1966:5). These extra cutaneous structures serve as

tactile organs for these nocturnal bottom-feeding fishes in

finding their prey. Similar filamentous processes are found

on the blind side of the head in certain sole-like pleuronectid

genera of the subfamily Rhombosoleinae, which also appear

to have nocturnal habits (Norman, 1934:20).

constant is the midlateral line, which extends from

the junction of the supraorbital canal with the

supraorbital commissures of the lateral-line system

of the head region and runs in the midaxis of the

body to the base of the caudal fin. In some species,

for example C. bilineatus, this line branches into

three at the base of the caudal fin, the middle running along the axis of the caudal fin, and the

other branches running parallel to the middle one

and extending onto the caudal fin. The next lateral

line of more or less constant occurrence on the ocular side in most of the species is the dorsolateral

line, which starts from the junction of the cephalodorsal canal with the supraoccipital commissure

of the head region and proceeds posteriorly in a

zig-zag manner along the dorsal edge of the abdominocaudal region, extending into the dorsal fin a

little distance behind the caudal base. The position

of the entrance of this line into the dorsal fin

varies in the different species and among individuals of the same species. The third and the least

constant lateral line of the ocular side is the ventrolateral line, which starts at the posterior end of

the base of the pelvic fin and runs posteriorly in

a zig-zag manner along the edge of the anal fin and

extends onto the anal fin a short distance before

the caudal. The place of entrance of this line onto

the anal fin also differs in different species and

even among individuals of the same species.

In some of the species of the Cynoglossinae the

ventrolateral line is well developed, but the presence or absence of this line should not be depended

upon for species differentiation, as this character

is highly variable. Norman (1926:300) had already

hinted at the unreliability of this character for

species or generic differentiation in cynoglossid

fishes and observed: "The number of lateral lines

on the ocular side may perhaps prove eventually

to be of doubtful value as a specific character.

Jordan and Starks (1960a: 240) have noted that in

Cynoglossus (Areliscus) interruptus the lower lateral line is broken at irregular intervals and often,

especially in smaller examples, it is entirely absent." I have verified this character in a number

of species of Cynoglossus and also in the case of

Paraplagusia guttata and find that there is considerable variation in the development of this lateral

line among individuals of the same species and

even of the same size.

As stated earlier, the midlateral line is the most
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

constant and well developed and is found in all

the species of the Cynoglossinae, while the dorsolateral, though present in all the species except C.

sinusarabici, is highly variable as regards its nature

and extent. It is, however, fully developed in all the

species where the ventrolateral is present.

THE SCALES

Along the lateral line in the Cynoglossinae the

pored scales usually have the perforating duct simple at its anterior opening, but in C. senegalensis,

C. lingua, and C. dubius it is different. In these

species the midlateral canal on the ocular side, instead of opening through simple pores on every

scale, opens by means of ducts into the adjoining

scale above or below. In C. senegalensis single ducts

alternate with two ducts, whereas in C. lingua

there are two ducts opening on one side, and in

C. dubius there are three openings on one side

followed by an opening on the opposite side.

The scales are generally ctenoid, the scales of the

lateral line having an imbricate arrangement with

the lateral-line canal in the middle. In all the

species with ctenoid scales the lateral-line scales are

provided with spinules above and below the canal.2

A good deal of variation in the nature of the scales

of the two sides, as well as those of the lateral line,

occurs among Cynoglossinae. When the scales are

ctenoid the spinules are invariably more strongly

developed on the ocular than on the blind side. In

some species the scales of the ocular side are ctenoid

and those of the blind side are cycloid, while in

certain others the scales of both sides are cycloid.

In some species the scales of the lateral line are

cycloid, while all those scales on the ocular side

independent of the lateral line are ctenoid (C.

lingua, C. senegalensis, C. bomeensis, C. browni, C.

bilineatus, C. lachneri, C. dispar, C. canariensis).

Regarding the nature of the scales of C. cadenati,

Chabanaud (1956) observed that the scales of the

upper side are ctenoid (at least in the young), including those of the lateral line (pleurogammiques),

and all the pores are simple. On the blind side

all the scales are ctenoid (except those of the head,

which are cycloid), but the scales toward the fin

region are considerably reduced; the marginal spin1

 In the Soleidae and Archiridae the scales of the lateral

line are much reduced, are enclosed in the skin, and are concealed by the two adjacent series.

ules are few, lying just above the surface or embedded in the skin. In the case of C. gilchristi Regan the scales and pores are as in C. cadenati, but

the scales in the region of the fins on the blind side

are more developed and the marginal spinules are

prominent. The rest of the species occurring off the

Atlantic coast of Africa, namely, C. senegalensis

Kaup, C. guinensis, C. canariensis (= C. lagoensis),

C. monodi, and C. browni, form a distinct group

characterized by the ctenoid condition (at least in

the young) of the upper scales except for the scales

on the lateral lines, which are cycloid; by the diverticulated opening of the lateral-line pores; and by

the cycloid condition of the scales of the blind side.

Thus, in the nature of its scales, C. cadenati differs

entirely from all the other species of Cynoglossus

occurring off the Atlantic coast of Africa, and it

should undoubtedly be placed in the group containing C. gilchristi, which is also characterized by

ctenoid scales (at least in the young) on both the

blind as well as the eyed side. Chabanaud (1956)

further concluded that in the genera Cynoglossus

and Paraplagusia the pleuragrammique formula

(number of lateral lines on the two sides of the

body) should be subordinated to the pholidologique

character (nature of scales).

Modifications of Scale with Age: Modifications in

the nature of the scales occur with age (size) in

certain species. For instance, in big specimens of C.

arel, which have mainly ctenoid scales including

those of the lateral line, a number of cycloid scales

are found in the lateral line toward the posterior

end, especially near the base of the caudal fin. The

scales that remain most consistently ctenoid are

those of the rows close to the dorsal and the anal

fins.

Dealing with modifications of scales among Cynoglossus, Chabanaud (1965) divided Cynoglossus into

two groups according to the nature of scales of the

upper side, namely, those with the entire upper

side with ctenoid scales and those with the lateralline scales cycloid and the rest of the scales ctenoid.

However, in the case of big specimens of C. arel,

which are characterized by ctenoid scales on the

upper side including those of the lateral line, a

number of cycloid scales occur in the lateral line.

Modification in the nature of the scales on the upper side is correlated with age, beginning from the

posterior end of the fish and at the base of the

caudal fin. The most conservative scales are in the
NUMBER 238 11

z^l^lfV

*mm*

FIGURE 2.—Diagramatic representation of lateral lines in Cynoglossus to show the variations in

the perforating ducts in the different species: a, C. senegalensis where the scales on the lateral

line are cycloid with single duct opening outside, alternately followed by two ducts on either

side; b, C. lingua where the scales on the lateral line are cycloid with two ducts opening on

either side in the adjoining scales; c, C. dubius where the scales on the lateral line as well as

the adjoining scales are cycloid with three ducts opening on one side followed by a duct

opening on the opposite side; d, Cynoglossus species where all the scales on the ocular side

are ctenoid with a single pore opening on every lateral line scale. (D = dorsal side, V = ventral side, C = caudal side, PO = pore opening, DI = diverticulum, CA = lateral line canal.)

rows closest to the dorsal and anal fins, the rest

gradually becoming cycloid with age. It is, therefore,

not generally possible to decide to which of the

two groups a specimen belongs unless juveniles or

immature specimens are examined.

T HE EPICRANIAL BONY SYSTEM

In the generalized genus Psettodes, the first spine

of the dorsal fin lies on the nape and well behind

the eyes, but in all other flatfishes the fin has ex-
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tended forward at least to above the eye and in

some even to the extremity of the snout. This forward extension of the dorsal in the Pleuronectoidea

is along the ridge of the supraoccipital and hence

along the bar formed by the union of the prefrontal and frontal on the blind side above the

upper eye, i.e., along the pseudomesial bar instead

of along the morphological median line. Above the

eye the fin is, therefore, not infrequently bent over

a little toward the blind side. In some genera, such

as Pleuronichthys, one or more rays of the anterior

part of the fin are virtually on the blind side of the

head.

The anterior extension of the dorsal fin onto

the head in flatfishes appears to have been achieved

either by (1) the anterior interneural spines themselves moving forward along the upper surface of

the cranium from a position in the region of the

supraoccipital to one on the frontal of the blind

sides as in Pleuronectes or (2) as in the case of the

Soleoidea, the first few interneural spines being inclinded forward, so that the first of these may actually occupy a horizontal position, thus carrying

the rays of the dorsal fin to the required place

(Norman, 1934:15). The interneural spines, inclined forward onto the cranium, are supported by

a hard bony system called the epicranial bony system (Ochiai, 1966:55).

In the case of the Soleoidea those anterior interneural spines that are inclined forward are supported by a special curved spinelike bone lying

nearly parallel to the cranium. This rod-shaped

bone is considered as the modified first interneural

spine and is called the erisma3 by Norman (1934:

15). Its base lies at a point between the ventral side

of the anterior end of the first neural spine and

the cranium, and its anterior end reaches to the

snout. In the Soleidae (except Heteromycteris

Kaup) and the Symphurinae the erisma is comparatively short and arched, its dorsal side near the

anterior end bearing two pterygiophores but no

interneural spines. In the specialized Heteromycteris

the erisma is long and thick, the anterior end being

strongly bent ventrally, supporting 15 interneural

spines dorsally and two long pterygiophores anteriorly (Ochiai, 1963:13).

In the Cynoglossinae the erisma is quite long and

'This bone is called "lame osseuse susbuccale" by Wu

(1932) and "axonoste libra" or "pseudo-axonoste" by

Chabanaud (1934b:284).

sickle shaped, supporting 10 interneural spines and

two pterygiophores dorsally. The anterior pterygiophore is specially well developed to form an axeshaped rostral cartilage (Wu, 1932:21) devoid of

any fin rays. This rostral cartilage in most of the

species is strong and flexible with 12 strong radially

arranged muscles on both sides. The erisma in the

Cynoglossinae is supported by one or two spinous

or sword-shaped bones, the pseudointerneural

spines. The anterior pseudointerneural spine is

sword shaped in most species, its base fixed lightly

on the frontal bone and the broader anterior portion extending forward onto the ventral side of

the erisma. In Symphurus the anterior pseudointerneural spine4

 is slender and spinous and its base

is attached to the supraoccipital bone.

The posterior interneural spine, which is spinous,

is present in most members of the Cynoglossinae

and the Symphurinae. In the Cynoglossinae it lies

between the cranial bones and the anterior pseudointerneural spine, while in the Symphurinae its

base is attached to the supraoccipital and it projects forward parallel to the anterointerneural

spine.

The Adaptive Significance of the Erisma: In

Pleuronectes a thin membranous bone is developed

on the middle axis of the cranium for supporting

the interneural spines that have moved onto the

head (Ochiai, 1963:56, fig. 14). This structure

strengthens the fin rays but prevents further invasion of the dorsal fin through the middle axis to

the anterior edge of the eye or snout region. In

most species of pleuronectids the dorsal fin, therefore, does not move onto the head portion beyond

the eye; if it does, it reaches to the area near the

nostril on the blind side. Those fin rays that reach

the snout region of the blind side are supported by

interneural spines on the prefrontal bone. In the

Soleoidea the erisma is separated from the cranium

and lies parallel to the middle axis of the head,

thereby enabling the dorsal fin to move easily along

the dorsal edge of the head as far as the snout

without being deflected to the blind side.

In the Soleidae the epicranial bony system is developed primarily to support the dorsal fin rays

on the head. In the process of specialization to a

bottom living habit and for fast swimming by

means of undulating movements, the vertical fins

* This bone is called "axonoste epicranier anterior" by

Chabanaud (1940:182).
NUMBER 238 13

have extended onto the head. The modification of

the pelvic fin, which is attached to the anal fin by

a membrane, is also a means of achieving efficient

undulating movements of the body. In the case of

Heteromycteris, the erisma extends anteriorly and

is bent ventrally to carry on its dorsal side the

interneural spines and the dorsal rays. In addition

to carrying those dorsal rays that have extended

forward to the head, the erisma has the secondary

function of strengthening the snout. In this case,

however, there is no rostral cartilage in the anterior portion of the erisma. In the case of Cynoglossus, the dorsal fin does not extend far forward

and the first pterygiophore does not bear any fin

ray but is modified into a rostral cartilage to give

support to the snout, thus enabling these fishes to

plough the sandy or muddy bottom to secure their

food, as well as to hide themselves by burying

wholly or partly in the sand. The poorly developed

sensory system, or lack of one, on the blind side of

the head and body of Cynoglossus, in contrast to

its development in the Soleidae, points to two distinct types of habit in these two groups of fishes.

While members of the Soleidae are mostly feeders

on the surface of the bottom by means of their

well-developed tactile organs on the blind side of

the head, the Cynoglossidae are adapted primarily

to ploughing the sand in search of prey and to feeding on organisms living below the surface. In Symphurus, although the erisma is not so well developed as in Cynoglossus or Paraplagusia, it is also

strengthened by the two pseudointerneural spines,

suggesting a similar habit of burrowing into the

bottom for feeding.

THE SPECIES GROUPS AND COMPLEXES AND

THEIR HYPOTHETICAL EVOLUTIONARY

RELATIONSHIPS

Taxonomically the genus Cynoglossus has several

lines of evolution representing six groups and 17

complexes.

The western periphery of the geographical range

of Cynoglossus, namely the tropical Atlantic coast

of Africa, is occupied by the most unspecialized

species closest to the ancestral stock. Toward the

Malay Archipelago are the more specialized and

rapidly evolving species complexes. The primitive

species or their immediate ancestors apparently

evolved in this center and were replaced subsequently by the more specialized species, such as

those of the cynoglossus, the carpenteri, and the

heterolepis groups.

The canariensis group, comprising the canariensis,

the browni, the bilineatus, and the attenuatus

complexes, is considered primitive and nearest to

the ancestral stock. These species are peripheral

in distribution (Figures 4, 5) and are mostly large

adult fishes reaching to about 400 mm. They exhibit

characters considered primitive in the process of

specialization toward a burrowing mode of life.

They generally have one or two lateral lines on the

blind side, 12 rays in the caudal fin, relatively big

eyes separated by a wide interorbital space, two

nostrils on the eyed side, large scales, and a rounded

snout. The possession of a lateral line on the blind

side is considered a primitive character for cynoglossid fishes feeding on organisms living below the

surface by digging or burrowing into sand or mud.

No useful purpose is likely to be served by having

lateral lines on the blind side and, therefore, they

are totally absent in the specialized forms. The

reduction in the number of caudal fin rays, reduced

size of scales, reduced size of eyes, scaly covering of

eyes, and loss of posterior slitlike nasal opening,

leaving only the anterior tubular nostril, are all

specializations for a burrowing mode of life.

The canariensis complex is the most primitive

complex of species within the canariensis group and

is characterized by a single lateral line on the blind

side. Its members are found at the extreme western

extremity of the generic range (Figure 4). The

bilineatus complex is the next evolved group. It

extends to the northern (Japan) and the southern

(Australia) periphery (Figure 5) and is characterized

by two lateral lines on the blind side.5 The attenuatus complex of species, occupying the east African

coast, the Seychelles, and the seas of India and

Pakistan (Figure 5), retains the primitive two

lateral lines on the blind side but shows a reduction

in the number of rays in the caudal fin; otherwise,

there are no structural changes from the bilineatus

complex. Cynoglossus browni is placed in a separate

•The bilineatus complex with two lateral lines on the

blind side is considered less primitive than the canariensis

complex, which has a single lateral line on the blind side

purely on a zoogeographical basis. It is probable that the

canariensis group evolved in west Africa from a form with

two lateral lines on the blind side and dropped off one

lateral line secondarily in the course of evolution (Carl L.

Hubbs, pers. comm.).
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

complex because of the absence of any lateral line

on the blind side. Morphologically, and in its

distribution, C. browni is closely related to the

canariensis complex and is probably evolved from

a canariensis-like ancestor through the loss of the

lateral line on the blind side.

The kopsi group, with small adult size members

attaining about 200 mm, is distinguished by the

lack of any lateral line on the blind side, large

eyes, either contiguous or separated by a very

narrow interorbital space, and a short obtusely

pointed snout, with the angle of the mouth situated

distinctly nearer to tip of snout than to branchial

opening.

The contiguous or subcontiguous nature of the

eyes is considered a very useful specialization in

fishes with a burrowing habit. In Cynoglossus two

lines of specialization of the eyes have taken place.

In the first case, where the eyes are of large size,

they have become contiguous or have come close

to one another, generally with the adipose membrane of the eyelid becoming thick and scaly,

holding the eyes more or less close together as a

compact structure, which gives sufficient protection

to them while ploughing or burrowing into the

sand. In the other case, which is more commonly

met with in the highly specialized species (heterolepis group), the eyes have become considerably

reduced in size, the interorbital width being the

same or even more than the diameter of the eye.

Included in the kopsi group are the kopsi, itinus.

ogilbyi, ecaudatus, and sealarki complexes. Of these,

the kopsi complex is considered the most primitive,

i.e., nearest to the canariensis group, the body scales

being somewhat larger than in the members of any

other complex (interlinear scale count: kopsi complex 7-12, ecaudatus complex 10-12, ogilbyi complex 8 or 11-14 and sealarki complex 10-17).

Reduction in the size of the body scales accompanies

specialization. The large imbricate scales on the

body are a disadvantage for the burrowing mode

of life of a fish. The itinus, ogilbyi, ecaudatus, and

sealarki complexes comprise closely related species

and have evolved into distinct complexes of species

mainly through gradual reduction of the size of the

body scales from the original kopsi stock that had

spread to different ecological nitches along the

coasts of Japan, Australia, and Africa. Today the

kopsi complex is restricted to an area from the

Indo-Australian Archipelago to southern Japan and

westward through India and Pakistan to Cargados

Garajos, Seychelles, and the Persian Gulf (Figure

7). The kopsi complex is characterized by two

nostrils on the ocular side, large body scales, ctenoid

on both sides, and 10 rays in the caudal fin.

In the African waters, two major lines of evolution of the early kopsi stock seem to have taken

place, one along the South African coast leading

to the sealarki complex of species and the other

along the north-east and West coasts of Africa

including the Red Sea and the Mediterranean to

the ecaudatus complex (Figure 7). The ecaudatus

complex is characterized by two nostrils on the

ocular side, comparatively small body scales with

10-12 interlinear rows, ctenoid on both sides, and

usually with 8 rays in the caudal fin, while the

members of the sealarki complex have only the

anterior tubular nostril, the posterior slitlike nostril

being absent, smaller body scales with 10-17 interlinear rows, ctenoid on the ocular, and partially

cycloid on the blind side, and 10 rays in the caudal

fin.

The stock that had spread from the Indo-Malayan

Archipelago southward to the Australian waters

differentiated into the ogilbyi complex of species

characterized by two nostrils on the ocular side,

smaller body scales with 8 or 11-14 interlinear

rows, ctenoid on the ocular side, and usually

cycloid on the blind side, and 8 rays in the caudal

fin. The itinus complex seems to have evolved in

the Japanese waters through the loss of the posterior

slitlike nasal opening, in all the other morphological

features itinus remaining similar to the kopsi

complex.

The arel group, with relatively large adult size

attaining about 400 mm, is a step forward in the

evolutionary history of the genus. The members of

this group are characterized by fairly large eyes but

separated by a narrow interorbital space, much

elongated body, and long obtusely pointed snout

with the angle of the mouth situated nearer to the

branchial opening than to tip of the snout. Its

range covers the whole of the Malay Archipelago

extending north to Japan and westward to India,

Pakistan, and the Persian Gulf (Figure 6). This

group has certain similarities to the canariensis and

bilineatus complexes, especially in the large adult

size of its members and the nature and position

of the eyes and large scales; therefore, it is considered to have evolved as an early offshoot from
NUMBER 238 15

CANARIENSIS

COMPLEX

C. BROWNI

BILINEATUS

COMPLEX

ATTENUATUS

COMPLEX

OCILBYI

COMPLEX'

KOPSI

COMPLEX '

C. ITiNUS

ECAUDATUS

COMPLEX

SEALARKI

COMPLEX

C. MONOPUS

CYNOGLOSSUS

COMPLEX

C PUNCT1CEPS

C.LIDA

CARPENTERI

COMPLEX

HETEROLEPIS

COMPLEX

CYNOQLO88U3 STOCK

FIGURE 3.—Dendogram showing the hypothetical evolutionary relationship of the species

complexes of Cynoglossus.

the main stock, which gave rise to the canariensis

group.

The cynoglossus group, consisting of fishes of

small adult size attaining only about 175 mm, is

characterized by relatively small eyes (pedunculate

in monopus complex) separated by a narrow interorbital space, a moderately elongated snout, but

with the angle of the mouth situated nearer to the

tip of the snout than to the branchial opening

(except lida complex), and relatively small scales on

the body, the interlinear scale rows being 7-21.

Its range covers the Malay Archipelago, northwest

Australia, the seas of India and Pakistan, and the

East Coast of Africa (Figure 8). In evolutionary
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

relationships, this group is considered closer to the

arel group, especially in the nature and position of

the eyes, the lateral-line system, and the caudal fin

ray count, than to the kopsi group. Hence it has

been considered as a recent offshoot from the stock

that gave rise to the arel group.

The carpenteri group, attaining relatively large

adult size (about 300 mm), is most probably derived

from an early cynoglossus-like ancestor through the

elongation of the body, the modified snout with the

shifting of the angle of the mouth to a point

distinctly nearer to the branchial opening than to

the tip of the snout, a reduction in the size of the

scales with an increased interlinear scale rows of

15-22, and the addition of a third lower lateral

line.

The heterolepis group is closely related to the

carpenteri group, differing mainly in having small

or minute eyes with a wide interorbital space. They

cluster mostly around the Malay Archipelago

(Figure 10) and are most probably evolved as an

early offshoot of the stock that gave rise to the

carpenteri group.

The species complexes have been erected on

Zoogeographical basis, and since characters such as

the loss of posterior slitlike nares occurred independently in widely separated areas and are

mainly adaptive, no separate key for their identification is given. However, a key for the identification

of the species has been provided.

Genus Cynoglossus Hamilton-Buchanan, 1822

Cynoglossus Hamilton-Buchanan, 1822:32, 365 [type-species:

Cynoglossus lingua Hamilton-Buchanan, monotypy.]

Cantoria Kaup, 1858:106 [type-species: Plagusia potous (not

potous of Cuvier) Cantor = Cantoria penanganensis Kaup,

monotypy.]

Arelia Kaup, 1858: 107 [type-species: Pleuronectes arel

Schneider = Arelia Schneider Kaup, tautonymy.]

Icania Kaup, 1858:109 [type species: Plagusia cynoglossus

Cantor = Icania cynoglossus Kaup, monotypy.]

Trulla Kaup, 1858:109 [type-species: Trulla cantori Kaup =

Plagusia trulla Cantor, tautonymy.]

Areliscus Jordan and Snyder, 1900:380 [type-species: Cynoglossus joyneri Giinther, monotypy.]

Cynoglossoides Bonde, 1922:23 [type-species: Cynoglossus

attenuatus Gilchrist, 1905, monotypy.]

Cynoglossoides [not Bonde, 1922] Smith, 1949:165 [typespecies: Cynoglossus ecaudatus Gilchrist, 1908, original

designation; preoccupied by Cynoglossoides Bonde 1922.]

Dollfusichthys Chabanaud, 1931:304 [type-species: Dollfusichthys sinusarabici Chabanaud, monotypy.]

Dexiourius Chabanaud, 1947b:443 [type-species: Cynoglossus

semilaevis Giinther 1873b, monotypy.]

Body lanceolate, eyes on left side, center of

migratory eye usually placed in advance of center

of fixed eye and usually anterior to its anterior

border, pectorals absent; only pelvic fin of blind

side present, with four rays, all inserted ventrally

in front of anal and connected to it by a membranous extension of its last ray. Dorsal, anal, and

caudal are confluent. The abdominocaudal region

of the ocular side always bears a mediolateral line

that, except in Cynoglossus sinusarabici, is accompanied by a marginodorsal line and in some species

by a third sensory line, the marginoventral. The

dorsolateral line generally runs in a zigzag or interrupted manner posteriorward and enters the dorsal

fin, the position of the entrance varying from species

to species and even among individuals. The ventrolateral line, when present, originates from the

posterior end of the base of the pelvic fin and also

runs in a zigzag or interrupted manner along the

base of the anal fin, entering the anal fin some

distance before the base of the caudal. On the

blind side the lateral lines are generally fewer in

number than on the ocular side; one or two or

often absent. On the ocular side of the head a

complex system of lines is present, which in its

essentials is invariable.

The scales are generally ctenoid on the ocular

side, ctenoid or cycloid on the blind side; the

scales on the mediolateral line are imbricate and

traversed by the lateral-line canal, the scales having

spinules above and below the pores of the lateral

line. On the ocular side the anterior nostril is

tubular, arising in front of the fixed eye a short

distance from the sublachrymal sulcus; the posterior

nostril when present is generally a simple opening

in the interorbital space when the eyes are separate,

or somewhat in front of the middle of the eyes when

the eyes are contiguous; on the blind side the

anterior nostril is generally tubular, arising above

the anterior half of the upper lip, the posterior

opening or slit generally lying at a level slightly

higher and above the posterior half of the upper

lip, the internarial space varying from two-thirds to

three-fourths of the distance between the posterior

nostril and the corner of the mouth opening. The

jaws are strongly asymmetrical; only the jaws of the

blind side are armed with teeth; the latter are needle

shaped, arranged on the dentary and the premaxil-
NUMBER 238 17

lary, forming a wide, short band with a meniscoidal

outline. The mouth is rather narrow, the snout

hooked and overhanging the mouth opening. The

lips are not fringed (cf. Paraplagusia). The gill

opening is narrow; the gill membranes are united,

free from isthmus. The urinary papilla is on the

eyed side and wholly joined to the first anal ray.

The ovarian duct opens into the posterior half of

the anus situated on the blind side.

The length of the neurocranium is less than its

height; the cerebral cavity is longer than the

rhinophthalmic part of the skull; the vault of

the cavity is complete; the sacculi are not

prominent.

Vertebral diapophyses are absent. Abdominal

vertebrae 9 or 10, rarely 11 or 12, caudal vertebrae

33 to 66. The closure of the abdominal haemal

arches is accomplished by a transverse bridge,

beyond which the two half-arches are prolonged

freely, on each side of the kidney.

The cleithrum is subangularly arched, the cuticular crest of the coracoid segment forming an

acutely angled apophysis at its base.

The epicranial bony system is well developed; the

erisma is angularly arched and prolonged anteriorly,

supporting a strong preoral cartilaginous plate.

Species of Cynoglossus were unknown to 18thcentury naturalists. The first species of Cynoglossus

known to science was Schneider's Pleuronectes arel

from Tranquebar, on the east coast of India. Two

years later Lac£pede (1802) described a species from

China and the East Indies under the name Achirus

bilineatus.

Hamilton-Buchanan's genus Cynoglossus was the

seventh in his order Apodes, consisting of "fishes

having the dorsal spine of bone and wanting ventral

fins." He characterized the genus as comprising

fishes "with both eyes on one side of the head and

with a flat body, formed for swimming on the side

opposite to the eye." Under his genus Cynoglossus,

Hamilton-Buchanan included only one species, C.

lingua. In the same work Hamilton-Buchanan

described another species, Achirus cynoglossus,

under his fourth order Thoracini, comprised of

fishes having the dorsal spine of bone and ventral

fin placed immediately under the pectorals. He

recognized, however, the close relationship of this

species to C. lingua.

In 1949, Chabanaud (1949a-d) published a series

of four papers on the Cynoglossidae of the Atlantic

coast of Africa. In the first of these he gave the

principal characteristics of the various genera of

the Cynoglossidae (1949a:60-64) and described C.

monodi from Dahomey and Sierra Leone (1949a:

65-66), giving a key to all the species, subspecies,

and morpha of Cynoglossus from that region.

Regarding the nature of the scales in C. monodi

he remarked (1949a: 66) that on the upper side all

the scales on the lateral line are cycloid; the pores

of the sensory canal are more or less distinctly

separated; the scales on the body are cycloid for

about the three quarters of the body length in the

type except for those that are placed near the

dorsal fin as well as in the ventral fin; those scales

are all ctenoid on the last quarter of the body in

the type. The scales on the blind side are cycloid.

He observed that by comparison with what exists

among the varied species of the Indo-Pacific, it is

reasonable to think that in some smaller (younger)

individuals all the scales on the upper side of the

body (nonpleurogrammiques) are ctenoid, while in

larger individuals these scales can become uniformly

cycloid. In the second paper in the series, Chabanaud (1949b) redescribed all the species of Cynoglossus and in the third and fourth (1949c,d) gave

a key and made certain remarks on the various

eastern Atlantic species. In the same year (1949e)

Chabanaud also described C. cleopatrides from the

Suez Canal.

In 1950 Chabanaud redescribed C. cleopatrides

and in 1951 published four important papers. In

the first of the series (1951a) he briefly discussed

the nature of the lateral line in C. senegalensis, C.

lagoensis, and C. kopsi. In the next (1951b) he

redescribed C. brachycephalus and described a form

from the Seychelles similar to C. brachycephalus,

but with three lateral lines in the abdominocaudal

region of the ocular side, which he tentatively

considered as a distinct species and provisionally

named it C. seychellensis. A form of C. brachycephalus from the Gulf of Suez, with a single lateral line

on the abdominocaudal region of the ocular side

(for which he had earlier erected a new genus and

species, Dollfusichthys sinusarabici), he considered

as yet another morphological variety of C. brachycephalus. He further observed that the morphological variety of a species is a matter of

geographical localization of its representatives. In

Indo-Malayan and (in a general way) Indian Ocean

species the upper lateral line is unbroken. To the
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

west of the Indian Peninsula, i.e., in the Persian

Gulf and the Red Sea but excluding the Gulf of

Suez, the upper lateral line is shortened not only

anteriorly, but also from its posterior end to the

point that in certain individuals, particularly those

that live in the Red Sea, the line in question

occupies no more than a minimum extent of the

middle part of the abdominocaudal region. Finally,

the same lateral line disappears entirely in the 22

specimens examined from the Suez Canal. The

Seychelles specimen with three lateral lines could

be an exceptional case because not far from its

place of capture the individuals found in the

Carajos belong to the two lateral-line types.

NOTE ON SYNONYMY.—As indicated by Norman

(1926), I find that the number and nature of

nostrils and the number of lateral lines on the

ocular side are of little value in generic differentiation. Cantoria is characterized by two nostrils on

the ocular side, both located above the upper lip;

Arelia by the presence of two nostrils, one tubular

on the upper lip before the eyes and the other

between the eyes; Trulla by the presence of only one

nostril in front of the lower eye; and Icania with

no conspicuous nostril at all. Giinther (1862) did

not consider any of Kaup's genera distinct and

grouped them all under Cynoglossus.

As an adaptation to a burrowing habit the

tubular narial opening is an advantage. It is,

therefore, invariably present in all such species,

whereas the posterior slitlike narial opening between

the eyes is sometimes absent. The posterior nares

are absent in the more highly evolved species like

the capensis complex of species (in Africa), C.

itinus (in Japan), and C. macrophthalmus (in

Australia); they are hidden by a thick fleshy covering as a protection against fine sand particles; they

are minute and invisible, unless examined through

a lens, in most of the other species.

Jordan and Starks (1906a) restricted the name

Cynoglossus to species with two lateral lines on

ocular side and placed those with three lateral

lines in Areliscus. Bonde (1922) considered forms

with two lateral lines on each side as Cynoglossoides

and commented (1925) on the utility of splitting

the original genus Cynoglossus into separate genera

and subgenera but observed that "if carried too far

it may lead to complications and an undue number

of monotypic genera." Further he observed that

"the lateral line which is the main character used

in splitting the genera is not always a constant

one." Smith (1949), obviously unaware of the

generic name Cynoglossoides proposed by Bonde,

included forms with two lateral lines on ocular

side and none on the blind side under his genus

Cynoglossoides, with C. ecaudatus as the typespecies. Chabanaud (1981) proposed Dollfusichthys

for a form characterized by a single lateral line on

the ocular side. The lateral lines, as has been

pointed out, are highly variable in number and

extant, and this cannot be considered as reliable

character for generic or even specific differentiation.

Chabanaud (1947c) proposed the genus Dexiourius for certain individuals of C. semilaevis from

China with a vestigial pelvic fin persisting on the

eyed side.

ZOOGEOGRAPHY AND EVOLUTION

DISTRIBUTION.—The geographical area of Cynoglossus comprises the eastern tropical Atlantic, the

eastern Mediterranean, the whole of the Indian

Ocean, including the Malay area in the east, the

Persian Gulf, the Gulf of Oman and the Red Sea,

the whole of the East Coast of Africa as far south

as the Cape of Good Hope in the west, the west

pacific from south China to south Japan, and the

whole of the periphery of the Australian continent.

The eastern and northern limit of Cynoglossus is

Tokyo at 35°40'N (C. interruptus); the southern

limit is the mouth of the Murray River, South

Australia, at 34°10/S (C. broadhursti). The western

limit is marked by the Canary Islands, about 30°N

(C. canariensis) in the Northern Hemisphere, and

Angola, about 10°S (C. canariensis) in the Southern

Hemisphere.

In spite of the much smaller spread into northern

and southern latitudes as compared to that of the

Soleidae,6 the distribution of Cynoglossus follows

a similar pattern of the Soleidae.

The most important conclusions that can be

drawn from an analysis of the distributional data

given in Table 2 are:

* The eastern limit of the Soleidae is the Galapagos Islands,

about 90°E (Aseraggodes herrei Scale), the western limit is

the tropical west African coast of Senegal, about 15*W

(Solea senegalensis Kaup), the northern limit is the North

Sea and west coast of Ireland, and the southern limit is

the Cape of Good Hope (Alagoa bay, etc.), about 34°S

(Solea bleekeri Boulenger and Astroglossus pectoralis (Kaup),

etc.).
NUMBER 238 19

TABLE 2.—General distribution of the species of Cynoglossus

(x = widespread distribution within area)

Species

West Medi- Red Arabia, India, Malay South Tai- New

Africa ter- Sea East Pak- Area China wan, Guinea,

ranean Africa istan Japan Australia

1. C.

2. C.

3. C.

4. C.

5. C.

6. C.

7. C.

8. C.

9. C.

10. C.

11. C.

12. C.

13. C.

14. C.

15. C.

16. C.

17. C.

18. C.

19. C.

20. C.

21. C.

22. C.

23. C.

24. C.

25. C.

26. C.

27. C.

28. C.

29. C.

30. C.

31. C.

32. C.

33. C.

34. C.

35. C.

36. C.

37. C.

38. C.

39. C.

40. C.

41. C.

42. C.

43. C.

44. C.

45. C.

46. C.

47. C.

48. C.

49. C.

canartensis

monodi

senegalensis

dubius

borneensis

browni

bilineatus

attenuatus

lachneri

dispar

kopsi

interruptus

joyneri

itinus

ogilbyi

maculipinnis

broadhursti

ecaudatus

cadenati

dollfusi

sinusarabicl

sealarki

microphthalmus ..

zanzibarensis

capensis

oral

robustus

lingua

cynoglossus

semifasciatus

macrostomus

monopus

puncticeps

durbanensis

gilchristi

lida

carpcnteri

acutirostris

marleyi

suyeni

heterolepis

feldmanni

abbreviatus

gracilis

semilaevis

macrolepis

kapuasensis

wandersi

tnacrophthalmus
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4 C CANARIENSIS

® C . MONODI

O C. SENEGALENSIS

A C. DUBIUS

* G . BORNEENSIS

• C BROWNI

FIGURE 4.—Distribution of canariensis-browni complex of species.

® C. BIUNEATUS

• CATTENUATUS

0 C . UCHNER1

CMSPAR

FIGURE 5.—Distribution of bilineatus-attenuatus complex of species.
NUMBER 238 21

60 90 120

40 4d

20*

# C. LIMGUA —

(§) C. AROi

| C. R0B0STUS

60° 90°

FIGURE 6.—Distribution of arel group of species.

r ; .•:.,..,»/. -I

B C . MACUUP1NM1S

BROADHURSTI

ECAUDATUS

SWUSARAHQ

SCD0LLFU 9

A C . JOTNERI

0CKOPSI

^ C DITERRUPTOS

FIGURE 7.—Distribution of kopsi, itinus, ogilbyi, and ecaudatus complex of species.
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

160*

® C . PUNCTICEPS

• C DURBANENSIS

* C . GILCHRIST1

FIGURE 8.—Distribution of puncticeps complex of species.

1. There is a decreasing number of species of

Cynoglossus as one progresses westward from the

Indo-Malayan region to the West Coast of Africa

through the coasts of India and Pakistan, the East

Coast of Africa, and the Red Sea.

2. The most primitive and generalized species

are found off the West Coast of Africa.

3. The most specialized forms are found in the

Indo-Malayan area.

Thus, the Malayan area, as the world's greatest

archipelago containing large areas with a depth of

less than 200 (Ekman, 1953), fufills all the conditions laid down by Matthew (1915) for a center

of dispersal of Cynoglossus. It has the largest

number of species, as well as the most progressive

forms, with the generalized species occurring at

the periphery of the range of the genus.

In considering the time of evolution of any group

of animals of which there is no fossil record, the

geological history of the areas it inhabits is essential.

At the beginning of the Tertiary, the Tethys Sea

covered large areas that today are dry lands, and

from the West Indies to the Pacific Ocean a rich

community of tropical animals populated the littoral zone of this area (Ekman, 1953:66). The

extinction of the tropical Tethys fauna in the

Mediterranean, according to Ekman, was mainly

due to the lowering of temperature in the late

Tertiary period when the northern elements of the

temperate fauna, which emigrated into the Mediterranean, formed the ancestral stock of the Mediterranean fauna of today. However, in the

Mediterranean, which is usually defined as a warmtemperate sea, it is the tropical forms that dominate

in the present-day fauna. These are mainly related

to the fauna of the warmer parts of the Atlantic

Ocean. There is, however, a northern boreal element as well in the present-day fauna but these

are Pleistocene immigrants (Kosswig, 1955). If it

were not for the Tertiary immigrants, the welladapted species from the north probably would

have occupied all the ecological niches in the

Mediterranean and thereby would have prevented

colonization by the later immigrants from the
NUMBER 238 23

4£_

C. ACUTJROSTRIS

*«-^-*r C. CARPENTERIV

FIGURE 9.—Distribution of carpenteri complex of species.

warmer parts of the Atlantic. Tha t there has not

been total occupation of all ecological niches in

the Mediterranean is evidenced by records of successful immigration of the Indo-Pacific forms into

it (Steinitz, 1927, 1929, Ben-Tuvia, 1966). Thes e

species coming from the Red Sea, the warmest and

saltiest sea (Marshall, 1952a,b), have passed through

the Suez Canal with its changing and extreme

ecological conditions and have established themselves in the eastern part of the Mediterranean.

T h e great plasticity exhibited by these species does

not lend any support to the view that their ancestors

died out in the Mediterranean at the end of the

Tertiary due to the climatic changes of that epoch;

indeed, they had several million years at their

disposal for adaptation to the cooling of the climate

that occurred then.

T h e occurrence today of Symphurus, the most

primitive genus of the Cynoglossidae, in both the

east and west Atlantic as well as off both the coasts

of America, suggests that it had most probably

evolved earlier than the Miocene when communication existed between the Indo-Pacific and the Atlantic through the Mediterranean and between the

American, Atlantic, and the Pacific through Central

America.

T h e present-day distribution of the Soleidae, including the broad soles, the Achiridae, extends

throughout the whole of the tropical Indian Ocean,

the Pacific, and the Atlantic. Soleid fishes are known

from the Eocene beds of Egypt (Woodward, 1910),

Java (Vorstman, 1927), and the Gulf Coast (Frizzel

and Dante, 1965), but no fossil records of any

cynoglossids are so far known. Among the cynoglossids, Symphurus has a much wider distribution,

coinciding with that of the Soleidae (including the

broad soles). From this distributional pattern it is

reasonable to conclude that Symphurus had evolved

much earlier than Cynoglossus, probably in the

Eocene, and at about the same time as the Soleidae,

later spreading westward as far as the Pacific coast

of America. Cynoglossus (including Paraplagusia),

the most specialized genus of flatfishes, seems to

have a more recent origin, probably not earlier than
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ISO*"

Em—to* . • .-.*/MARSHALL

COCO*-K*%klH«

CGRACIUS

C. ABBREVIATE

C. SEMILAEVIS

*> C. HETEROLEPSIS

Z. C. KAPUASENSIS

O C MACROPHTHALMUS

• C. FELDMANNI

• C. WANDERSI

i t C. MICROLEPS1S

FIGURE 10.—Distribution of heterolepis group of species.

DISTRIBUTION OF CYNOGLOSSIDAE

= PARAPLAGUSIA

= CTNOGLOSSUS

= SYMPHURUS

FIGURE 11.—Distribution of Cynoglossidae: Symphurus, Cynoglossus, and Paraplagusia.
NUMBER 238 25

Pliocene. The occurrence today of closely related

species of Cynoglossus—the canariensis and the

ecaudatus groups—off tropical west African coasts

on the one hand and in the Indo-Pacific on the

other is very significant in this connection (Figures

4,7).

GEOLOGICAL HISTORY OF THE RED SEA.—Geologically the Red Sea is relatively young, the main

physical features being completed only during the

Pliocene. The Red Sea came into being as a result

of faulting of Eocene strata and the filling of this

great rift with water from the Tethys Sea to the

north of it. During Miocene times this sea extended

southward as far as southern Egypt. This no doubt

was an extension of the Proto-Mediterranean for

its fossils are mainly of Mediterranean affinities

(Davies, 1934). The upper Miocene movements

caused the elevation of the region and a period

of continental conditions prevailed over the gulf

of Suez during late upper Miocene and lower Pliocene (Said, 1962). A connection between the Red

Sea and the Indian Ocean seems to have been established during the latter half of the middle

Pliocene, when a phase of subsidence took place

and the Red Sea proper was flooded by the Indian

Ocean through a break at the Bab-el-Mandeb

Strait (Fox, 1926). Marine Pliocene deposits are

noted as far south as latitude 28°30'N, to the south

of which there is definite proof of marine invasion

by Indo-Pacific marine Pliocene fauna (Said, 1962:

194). To the north of this latitude and caused by

the same tectonic movements, considerable sinking

of the Gulf of Suez also took place during the middle Pliocene, for the deposits of these periods, especially along the Sinai Gulf coastal strip, are in

places more than 1500 m in thickness (Said, 1962).

In the north and east the upper Pliocene movements must have been strong enough to produce

flooding of the Isthmus of Suez and the free mixing

of water from the Mediterranean and the Gulf of

Suez, but it is uncertain when the Gulf of Suez

became cut off from the Mediterranean (Marshall,

1952a).

This would seem to be the generally accepted

geological history of the Red Sea, but Sewell (1948)

considered the possibility of the emergence of the

shallow sill at the southern end of the Red Sea

during the Glacial periods and the reduction of the

Red Sea to an inland lake (Marshall, 1952a). It is

probable that the Gulf of Suez was also cut off from

the Mediterranean during the glacial periods. An

excellent recent account of the geological history

of the Red Sea has been given by Botros (1971).

DISPERSAL DURING THE PLIOCENE TO RECENT.—

The earliest stock of Cynoglossus, namely the

canariensis group, seems to have spread from the

Malay area westward through the seas of India

and Pakistan to the Arabian coast and thence

through the Red Sea to the Mediterranean, presumably during the upper Pliocene. It must then

have spread northward through the Chinese Sea

to south Japan and through Indonesian waters

southward to Australia.

The kopsi group (consisting of the itinus, ogilbyi,

ecaudatus, and sealarki complexes), which is the

nearest to the canariensis group, also seems to have

spread to the Mediterranean along the same route

at a later period, but not later than early Pleistocene. Thus, C. cadenati is today found off tropical

west Africa, while the closely related species C.

dollfusi and C. sinusarabici occur in the Suez Canal,

the eastern Mediterranean, and the Red Sea (Figure 7).

The freshwater discharge of the Nile into the

Mediterranean must have formed a barrier to the

westward spread of many of the Indo-Pacific fishes

(Ekman, 1953:88). Cynoglossus, however, are

bottom-living euryhaline fishes, and the freshwater

discharge of the Nile and also the shallow strait

that probably existed at the present-day Isthmus of

Suez connecting the Red sea and the Mediterranean

(Tortonese, 1964:103) would not have formed a

barrier for their migration to the Mediterranean

and their establishment there.

It is not clear whether the present-day Mediterranean fishes have persisted there throughout the

Quaternary period. As stated above, it seems more

probable that the Tethys fauna died out as a result

of the lowering of temperature or changes in salinity in the Mediterranean during the glacial periods

of the Pleistocene. The Isthmus of Suez must have

been exposed, leaving probably only brackish water

lakes along its course. This separation of the Mediterranean from the Red Sea ended in 1869 when,

with the opening of the Suez Canal, fresh opportunities were afforded to the Indo-Pacific fishes to

enter into the Mediterranean. The record of C.

sinusarabici from Israel (Ben-Tuvia, 1966) indicates

its recent spread there.
2 6 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species

1. 12 rays in caudal fin 2

Less than 12 rays in caudal fin 8

2. Lateral line on blind side 3

No lateral line on blind side (west Africa) C. browni

3. 1 lateral line on blind side 4

2 lateral lines on blind side (Indo-West Pacific except Africa and Red Sea) C. bilineatus

4. Cycloid scales on ocular side 5

Ctenoid scales on ocular side 6

5. 17-19 scales between middle and upper lateral lines (west coast of India) C. dubiut

13-14 scales between middle and upper lateral lines (west Africa) C. tnonodi

6. 11-12 scales between middle and upper lateral lines (west Africa) C. canarietuis

More than 12 scales between middle and upper lateral lines 7

7. 16—18 scales between middle and upper lateral lines (west Africa) C. senegalensis

18-20 scales between middle and upper lateral lines (Borneo and Thailand) C. borneensis

8. 10 rays in the caudal fin 9

Less than 10 (usually 8) rays in caudal fin 39

9. 2 lateral lines on blind side 10

No lateral line on blind side 12

10. 10-11 scales between lateral lines on ocular side (south Africa) C. attenuatut

More than 11 scales between lateral lines 11

11. 16-18 scales between lateral lines (Mozambique coast and Oman) C. lachneri

18-2.0 scales between lateral lines (India and Pakistan) C. dispar

12. A single nostril on ocular side IS

2 nostrils on ocular side : 17

13. Ctenoid scales on blind side 14

Cycloid scales on blind side 15

14. 13-15 scales between upper and middle lateral lines (Zanzibar and Kenya) ... C. xanxibarensis

17-18 scales between upper and middle lateral line (Queensland) C. macrophthalmus

15. 10-11 scales between upper and middle lateral lines (Saya de malha) C. sealarki

More than 11 scales between lateral lines 16

16. 12 scales between lateral lines (Natal) C. microphthalmus

15-17 scales between lateral lines (Saldanha Bay, Natal) C. capensis

17. Both nostrils situated before eyes, eyes pendunculate (Malay Archipelago, Hong Kong, and

Bay of Bengal) C. monopus

The posterior nostril situated in interorbital space or between anterior part of eyes, eyes

nonpedunculate 18

18. Cycloid scales on blind side 19

. Ctenoid scales on blind side 25

19. 2 lateral lines on ocular side 20

3 lateral lines on ocular side 23

.20. 11 or more scales between lateral lines 21

Less than 11 scales between lateral lines 22

21. Corner of mouth nearer to gill opening than to end of snout; 11-12 scales between lateral

lines (Malay Archipelago, Hong Kong, Bay of Bengal) C. lingua

Corner of mouth nearer to snout than to gill opening; 12-14 scales between lateral lines

(Western and South Australia) C. broadhursH

22. 10 scales between lateral lines (south China to Japan) C. robustus

7-9 scales between lateral lines (Malay Archipelago and India) C. arel

23. Snout acutely pointed; angle of mouth distinctly nearer gill opening than to end of snout;

less than 20 scales between upper and middle lateral lines 24

Snout rounded; angle of mouth nearer tip of snout than gill opening; more than 20 (20-25)

scales between upper and middle lateral lines (China) C. semUaevis

24. Snout about 33 percent of head; 15-19 scales between middle and upper lateral line (coasts

of India 10 Persia) C. carpenter*

Snout longer, 43 percent of head; 18-20 scales between middle and upper lateral line (Gulf

o f Aden ) ; C. acutirostri,

25. Usually two lateral lines on ocular side 25
NUMBER 238 27

Usually three lateral lines on ocular side 32

26. Angle of mouth distinctly nearer to gill opening than to end of snout (Malay Archipelago

and Philippines, India and Africa) C. lida

Angle of mouth nearer to end of snout than to gill opening 27

27. Cleft of mouth extending far back posterior margin of fixed eye (India) C. macrostomus

Cleft of mouth extending only middle or about posterior border of fixed eye 28

28. Eyes contiguous (Malay Archipelago, Philippines, Taiwan, India, Cargados Garajos, and

Persian Gulf) C. kopsi

Eyes not contiguous 29

29. 14 of fewer scales between lateral lines SO

More than 14 (16-19) scales between lateral lines (Malay Archipelago to Philippines, Taiwan,

and India) C. puncticeps

30. Body slender, about 20 percent of SL, 11-12 scales between lateral lines (west Africa)

C. cadenati

Body deep, about 26 to 27 percent of SL, 10-14 scales between lateral lines 31

31. Snout rounded and short about 27 percent of head, 12 (11-14) scales between lateral lines

(east coast of India) C. semifasciatus

Snout somewhat pointed, longer, about 32 percent of head, 13 (12-14) scales between lateral

lines (Malay Archipelago, Philippines, and India) C. cynoglotsus

32. Eyes contiguous ..32

Eyes not contiguous 34

33. Angle of mouth nearer to gill opening than to tip of snout; 19-22 scales between upper

and middle lateral lines (Philippines, Celebes, and Timor) C. suyeni

Angle of mouth nearer to tip of snout than to gill opening; 11-12 scales between upper and

middle lateral lines (Japan) C. mterruptus

34. Less than 20 scales between upper and middle lateral lines 35

More than 20 scales between upper and middle lateral lines 38

35. Corner of mouth nearer to tip of snout than to gill opening; 11-12 scales between lateral

lines (south China and Japan) C. joyneri

Corner of mouth nearer to gill opening than to tip of snout 36

36. Interorbital space narrow, less than half eye diameter; 18-19 scales between upper and

middle lateral lines (Durban to Delagoa Bay) C. marleyi

Interorbital space wide, about the same as eye diameter 37

37. 15-16 scales between lateral lines (New Guinea and Western Australia) C. heterolepis

17-18 scales between lateral lines (Borneo and Sumatra) C. feldtnanni

38. Corner of mouth nearer to tip of snout than to gill opening; interorbital space much less

than eye diameter; 24 scales between upper and middle lateral lines (Sumatra and Borneo)

C. waandrrsi

Corner of mouth nearer to gill opening than to tip of snout; interorbital space much wider

than eye diameter; 25 scales between upper and middle lateral lines (West Borneo in

fresh waters) C. kapuasensis

39. 1 nostril on eyed side (Japan and Hong Kong) C. itinus

2 nostrils on eyed side 40

40. Eye contiguous 41

Eyes not contiguous 43

41. 1 lateral line on ocular side (Red Sea, Suez Canal, and east Mediterranean) . C. simuarabici

2 or 3 lateral lines on ocular side 42

42. Usually 2 lateral lines on ocular side (east Africa and Seycheles) C. ecaudatus

Usually 3 lateral lines on ocular side (Suez Canal) C. dollfusi

43. Cycloid scales on blind side (south Queensland) C. oplbyi

Ctenoid scales on blind side 44

44. Usually 2 lateral lines on ocular side 45

Usually 3 lateral lines on ocular side 47

45. 11-12 scales between lateral lines (northwest, north, and east Australia) C. maculipmnis

More than 12 scales between lateral lines 46

46. 14-15 scales between lateral lines (Natal to Madagascar) C. gilchristi

18-21 scales between lateral lines (Natal to Zanzibar) C. durbanensis

47. Body deep, more than 20 (24) percent of SL, 18-23 scales between upper and middle lateral

lines (south China through Taiwan to south Japan) C. abbreviatus
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Body slender, about 20 percent of SL 48

48. Upper eye distinctly in advance of the lower (Thailand, Cambodia, Borneo, and Sumatra, in

fresh waters) C. microlepis

Upper eye not in advance of the lower (south China through Taiwan to Korea)

C. gracilis

The canariensis group

Four species complexes, namely the canariensis

complex, the browni complex, the bilineatus complex, and the attenuatus complex, are here included

in what may be called the canariensis group, canariensis being closest to the evolutionary stem of the

group and probably of the entire genus. This group

(with the exception of C. browni) is characterized

by one or two lateral lines on the blind side, two

or three lateral lines on the ocular side, 12 rays in

the caudal fin (except the attenuatus complex),

large eyes separated by a wide interorbital space,

two nostrils on the eyed side, large scales, and

rounded snout.

The members of the canariensis group are the

most morphologically generalized species of the

genus, the presence of lateral lines on the blind

side being considered a primitive character in

Cynoglossus. The reduction in the number of caudal fin rays, the small or reduced size of the eyes,

the type of scales, and the atrophy or reduction in

the number of nostrils are all considered specializations for a burrowing habit.

The canariensis complex

Included in this complex are C. canariensis, C.

monodi, C. senegalensis, C. dubius, and C. borneensis. These are large species characterized by 12 rays

in the caudal fin and one lateral line on the blind

side. Their center of distribution is in west Africa,

the western extremity of the range of the genus;

apparently they are a very old group and probably

the earliest stock to be given off from the evolutionary center of the genus, namely the Malay Archipelago.

1. Cynoglossus canariensis Steindachner

FIGURE 12; PLATE 1

Cynoglossus canariensis Steindachner, 1882:13, pi. 2: fig. 2

[type-locality: Bane d"Argain, Canary Is.].—Chabanaud and

Monod, 1926:284.—Fowler, 1936:526, fig. 249.—Chabanaud,

1949b:203.—Cadenat, 1960:1383.

Cynoglossus lagoensis Regan, 1915:129 [type-locality: Lagos].—

Fowler, 1936:526.—Chabanaud, 1949b:207.

DESCRIPTION.—Based on 10 specimens 173.0-367.0

mm SL, including the lectotype and paralectotypes

of C. lagoensis.

Depths of body 21.39-26.45 (M = 24.60) and

length of head 20.23-24.19 (M = 21.51) percent of

standard length. Diameter of eye 6.29-8.57 (M =

7.13), interorbital width 4.29-8.0 (M = 6.13) percent of length of head. Two nostrils on ocular

side, anterior nostril of eyed side tubular, on upper

lip in front of lower eye, posterior nostril simple,

in anterior half of interorbital space. Snout pointed,

30.67-34.29 (M = 32.55) percent of length of head,

rostral hook short and hardly reaching to vertical

through front border of upper eye. Maxillary extending to well beyond fixed eye; angle of mouth

extending to beyond vertical from posterior border

of fixed eye, nearer to tip of snout than to branchial

opening; snout to angle of mouth 46.58-52.86

(M = 49.33), angle of mouth to branchial opening

50.67-57.33 (M = 55.08) percent of length of head.

Scales: On eyed side (except on lateral lines)

ctenoid in anterior part, cycloid posteriorly; scales

on blind side and on lateral lines of eyed side

cycloid.

Lateral-Line System: 3 lateral lines on ocular

side, midlateral line with 76-88 scales. 10-13 (M =

12) scales between upper and middle lateral lines.

One lateral line on blind side.

Interlinear scale rows 10 11 12 13

Frequencies 1 0 4 1

Fins: Dorsal with 125 and anal with 99 rays,

caudal 12 in 3 specimens (radiographs).

Vertebrae: 57-59, comprising 9 abdominal and

48-50 caudal elements in 3 specimens (radiographs).

Coloration: Upper side uniformly brownish, lower

rather whitish in preserved specimens.

Size: Largest specimen examined, 546.0 (510 + 36)

mm, is from Angola collected by the Discovery.

BMNH 1935.5.11.228.

DISTRIBUTION.—West Africa: Canary Islands,

Senegal to Angola.
NUMBER 2S8 29

Icm

OV

FIGURE 12.—C. canariensis: a, outline drawing of lateral view of ocular side of a specimen

(BMNH 1935.5.11.228) from Angola; b, details of anal region on blind side of lectotype (BMNH

1914.11.2.72). (A = anus, OV = ovarian orifice.)

DIAGNOSIS AND AFFINITIES.—Cynoglossus canariensis is closely related to C. monodi and C. senegalensis. It is, however, distinguishable by its relatively

larger scales, the interlinear scale count being

11-13 (12-14 in C. monodi and 17-18 in C. senegalensis), and the nature of body scales, ctenoid on

ocular side, cycloid on blind side (C. canariensis

and C. senegalensis), and mostly cycloid on both

sides (C. monodi).

NOTE ON SYNONYMY.—Steindachner's (1882:13)

description of C. canariensis was based on a single

specimen, 280 mm in TL, from the Canary Islands.

His fish had three lateral lines on the ocular side

and one on the blind side. Because of the excellence

of his description and illustration there has been

little confusion in the identity of his species. However, Regan (1915:129) described C. lagoensis on

the basis of three specimens, 390-512 mm TL from

Lagos and Angola, and related it to "C. canariensis

Steindachner in which the cleft of the mouth ends

below the middle of the eye, the head is smaller,

the scales are more numerous." I have examined

the lectotype and the paralectotypes of C. lagoensis

in London and found them referable to C. canariensis. Comparison of several specimens from west

Africa has shown that the differences noted by Regan are merely intraspecific variation.

TYPE SPECIMENS.—Holotype of C. canariensis,

NHV 47-773, 252 mm SL, 259 mm TL; Canary

Islands, Africa (pers. comm., Dr. Paul Kahsbauer).

BMNH 1914.11.2.72, 356 mm SL, lectotype of C.

lagoensis, designated by Chabanaud (1949b:207),
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lagos, Cadman. Another specimen, BMNH 1914.-

11.2.71, 365 mm SL, paralectotype of C. lagoensis,

designated by Chabanaud, Lagos, Cadman, and a

further specimen, BMNH 1935.5.11.288, paralectotype of C. lagoensis, designated by Chabanaud,

Angola, Discovery.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH

19353.11.229. Angola, coll. Discovery. 2, BMNH

1962.6.18.135.36, Atlantide Sta 86, coll. N. Nielson. 1,

BMNH 1962.6.18.137, Atlantide, 102.5'34'N, 4°50'E, coll. N.

Nielson. 1, ZMA 108.008, W Africa, Cape Bianco, coll.

Vermeulen, 1906. 5, USNM 188435, off Lagos, Nigeria, coll.

A. Longhurst. 1, CU, Ghana Teshis, Atlantic Drive, coll.

Amegah, 26.1.1962. 1, CU, Ghana, Amegah, 14.8.1961.

2. Cynoglossus monodi Chabanaud

FIGURE 13; PLATE 1

Cynoglossus monodi Chabanaud, 1949a:65 [type-locality:

Dahomey and Sierra Leone, west Africa].—Cadenat,

1960:1383 [Lagos, Nigeria; Chorkor, Accra, Ghana].

DESCRIPTION.—Based on 7 specimens, 281.0-327.0

mm SL, including the holotype of C. monodi.

Depth of body 20.00-23.85 (M = 21.46) and

length of head 19.65-22.0 (M = 20.16) percent of

standard length. Diameter of eye 6.25-7.08 (M =

6.50), interorbital width rather broad, 5.65-8.00

(M = 6.88) percent of length of head. Anterior

nostril of eyed side tubular, on upper lip in front

of lower eye, posterior nostril simple, in anterior

half of interorbital space. Snout narrowly rounded,

38.71-44.25 (M = 42.32) percent of length of head,

rostral hook rather short, stopping just in front of

vertical through front border of fixed eye. Maxillary extending to well beyond fixed eye; angle of

mouth extending to right behind vertical from

posterior margin of fixed eye, nearer to branchial

opening than to tip of snout; snout to angle of

mouth 52.42-58.04 (55.46), angle of mouth to branchial opening 38.52-46.77 (M = 44.05) percent of

length of head.

Scales: Cycloid on both sides, except on last

quarter of body on upper side, where they are

ctenoid.

Lateral-Line System: Two lateral lines on ocular

side, midlateral line with 85-96 scales, 12-14 scales

between middle and upper lateral lines. One lateral

line on blind side.

Inter linear scale rows

Frequencies

12 IS 14

1 2 8

Fins: Dorsal with 125-131 (M - 127) rays, anal

with 99-105 (M = 101) rays, caudal 12 in 3 specimens radiographs).

Vertebrae: 57-60 comprising 9 abdominal and

48-51 caudal elements in 3 specimens (radiographs).

Coloration: Upper side is light brown and the

lower rather whitish in preserved specimens.

Size: Largest specimen examined, USNM

193912. 352.0 (327 + 25) mm, is from Nigeria, off

Lagos.

DISTRIBUTION.—West Africa: Dahomey, Sierra

Lione, Nigeria, Accra, Chana, and Lagos.

DIAGNOSIS AND AFFINITIES.—Cynoglossus monodi

is closely related to both C. canariensis and C.

senegalensis, but it can be readily distinguished by

the cycloid nature of the scales on its ocular side

(ctenoid in C. canariensis and C. senegalensis).

FIGURE 13.—Outline drawing of C. monodi, paratype (BMNH 1949.4.30.4)

from Great Popo, west Africa.
NUMBER 238

TYPE SPECIMENS.—Chabanaud (I949a:65) described C. monodi on the basis of 3 specimens,

241-349 mm in TL, from Dahomey and Sierra

Leone. I have succeeded in locating only 2 specimens, the holotype, MNHP 49-18, 312 mm SL, and

one paratype, BMNH 1949.4.30.4, 237 mm SL. The

third specimen from Sierra Leone is likely to be in

the Paris Museum but is not traceable at the

moment.

OTHER MATERIAL EXAMINED.—3 specimens, USNM 188434,

Nigeria off Lagos, coll. A. Longhurst, 1962. 2. USNM

193910-12, Liberia, off St. Paul River, 1^-7 fms, coll. G. C.

Miller, 29 Dec 1952.

3. Cynoglossus senegalensis (Kaup)

FIGURE 14; PLATE 2

Arelia senegalensis Kaup, 1858:108 [type-locality: west Africa].

Plagusia (Arelia) senegalensis.—Dumeril, 1858:264 [Senegal].

Cynoglossus senegalensis.—Giinther, 1862:502.—Capello,

1872:86, [Bissau, west Africa].—Rochebrune, 1882:116

[Goree, Dakar, Guet N'Dar, Barbarie].—Buttikofer.

1890:480.—Steindachner, 1894:50 [Messurad River mouth,

Liberia].—Pellegrin, 1913:153 [Lagune de Porto-Novo,

Dahomey]; 1914:78 [Lagune de Porto-Novo, Dahomey].—

Fowler, 1936:527,1261.

Cynoglossus (Arelia) senegalensis.—Steindachner, 1870:977

[St. Louis].

Cynoglossus senegalensis senegalensis Chabanaud. 1949b:204

[morpha typica, two lateral lines on ocular side, one on

blind side]; 1949d:518.

Cynoglossus senegalensis simulator Chabanaud, 1949b:205

[Dakar].

Plagusia bilineata [not Lacepede].—Dumeril, 1858:246 [Isle

of Prince, Portuguese West Africa].

Cynoglossus goreensis Steindachner, 1882:12, pi. 1: fig. 2

[Goree].—Osorio, 1894:134 [Benguella, Angola]; 1898:199

[Angola].—Pellegrin, 1905:35 [Guet N' Dara, Dakar];

1913:152 [Konakry, Guinea]; 1914:79 [Guet N' Dar, Dakar;

Konakry].—Regan, 1915:129 [Lagos].—Fowler. 1918a:249

[Cape Palmas, Liberia], 268 [Gaboon].—Chabanaud and

Monod, 1926:284 [Cape Blanco].—Fowler, 1936:526 [Liberia

and Gaboon].—Chabanaud, 1949d:519.

Cynoglossus guineensis Osorio, 1909:104, pi. 1: fig. 2 [typelocality: Bolama, Portuguese Guinea].—Chabanaud,

1949b:202 [Dahomey and Togo]; 1949d:519.—Cadenat,

1960:1383 [Lagos, Nigeria, Jui, and region of Freetown,

Sierra Leone].

DESCRIPTION.—Based on 11 specimens, 117.5 mm375.0 mm SL, including the holotype of C. senegalensis simulator.

Depth of body 20.38-24.79 (23.13), length of head

18.63-22.13 (M = 20.26) percent of standard length.

Diameter of eye 6.08-9.68 (8.14), interorbital width

5.98-9.86 (M = 7.50) percent of length of head. Two

nostrils on ocular side, anterior tubular on upper

lip, the tube not extending to fixed eye, the posterior in the anterior half of interorbital space.

Snout prominent, broadly rounded, 24.62-39.32

(M = 33.84) percent of length of head, rostral hook

rather short, extending only to front of anterior

nostril. Maxillary extending beyond fixed eye; angle of mouth extending to just beyond vertical from

posterior border of fixed eye, slightly nearer to tip

of snout than to branchial opening; snout to angle

of mouth 47.30-54.29 (M = 50.04), angle of mouth

to branchial opening 47.14-56.45 (52.07) percent of

head.

Scales: Ctenoid on ocular side, except those on

lateral lines; scales of blind side and of lateral lines

on ocular side cycloid.

Lateral-Line System: Two or three lateral lines

on ocular side, midlateral line with 89-108 scales;

17-18 scales between upper and middle lateral lines.

One lateral line on blind side. The midlateral line

on ocular side instead of opening through simple

pores on every scale opens by means of simple ducts

into the adjoining scale, generally with a single

I cm

FIGURE 14.—Outline drawing of C. senegalensis (BMNH 18963331)

from Igowe River, Lambarene.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

duct on one side followed by two ducts on the

other side (Figure 2).

Interlinear scale rows 17 18

Frequencies 6 8

Fins: Dorsal with 119-125 (M = 121) rays, anal

with 93-99 (M = 96) rays, caudal 12 in 7 specimens

(radiographs).

Vertebrae: 56-58 comprising 9 abdominal and

47-49 caudal elements in 7 specimens (radiographs).

Coloration: Upper side is dark brown in alcohol

and blackish or greenish in formalin, the blind side

is white in preserved specimens.

Size: The largest specimen examined, 397 (375

+ 22) mm, holotype of Cynoglossus senegalensis

simulator Chabanaud, is from west Africa.

DISTRIBUTION.—West Africa: Liberia to Lagos.

DIAGNOSIS AND AFFINITIES.—Cynoglossus senegalensis is closely allied to C. canariensis and C.

monodi. These three form a fairly well-defined

complex of species, but C. senegalensis and C.

canariensis have ctenoid scales on ocular side (cycloid scales in C. monodi). Cynoglossus senegalensis

is, however, easily distinguished by its smaller body

scales, the interlinear scale count being 16-18 cf 10-

14 in C. canariensis, 12-14 in C. monodi.

NOTE ON SYNONYMY.—Kaup's (1858:101) description of Arelia senegalensis was meager and was

based on a single specimen, 555 mm in TL from

west Africa. A very salient feature in his description was the presence of two lateral lines on the

ocular side. Steindachner had the same species before him when he described C. goreensis on the

basis of a specimen, 600 mm in TL, from Goree and

characterized it by having three lateral lines on the

ocular side. Chabanaud (l949b:203) synonymized

it with C. senegalensis and I concur with him since

an additional lateral line is of no significance.

Osorio (1909:104) gave the name C. guineensis to

a specimen from Guinea on the ground that it

differed from C. senegalensis by its slightly smaller

scales and somewhat deeper body. Comparison of

numerous specimens from west Africa has convinced me that these differences are only intraspecific variation and I have, therefore, synonymized

C. guineensis with C. senegalensis. It may be

pointed out, however, that Osorio's description is

defective in that he mentions only 16 scales between the upper and middle lateral lines, whereas

in the illustration 18 rows are indicated.

Chabanaud (1949b:205) described C. senegalensis

morpha simulator based on a single specimen, 410

mm in TL from Dakar, and distinguished it from

C. s. senegalensis morpha typica by its possession of

an additional lateral line on the ocular side. Since

the development of an additional lateral line is

not considered of any significance in the differentiation of species in Cynoglossus, I have considered

the two species conspecific. I have examined the

holotype of C. s. simulator and found it to be

identical to specimens of C. senegalensis.

TYPE SPECIMENS.—The type specimen of C. senegalensis is not available in Berlin (pers. comm., C.

Karrer) and is most probably not extant. MHNP

49.22, 375 mm SL, holotype of C. senegalensis simulator, Nigeria, off Lagos, coll. A. Longhurst. NHV

3-776, 550 mm SL, 590 mm TL, holotype of C.

goreensis, Goree, Dakar, Senegal (Dr. Paul Kahsbauer examined the type on my behalf).

OTHER MATERIAL EXAMINED.—3 specimens, USNM 188433,

Nigeria, off Lagos, coll. A. Longhurst. 1, USNM 193653,

Mesurado R. beach, Liberia, coll. G. C. Miller. 1, USNM

193881. Bushrod Is., Liberia, coll. G. C. Miller. 1, UMMZ

202912, off St. Paul R., Liberia (4^-7 fms). 1, USNM 247424,

Nigeria, off Lagos. 2, CU, Volta R., Ghana, coll. Bane and

Richards, 23.7.1951. 3, CU, Labadi, Atlantic drainage,

Ghana, coll. Amegah, 26.1.62. 3, CU, Teshi, Ghana, coll.

Amegah, 26.1.62. 3, BMNH 1862.1.22.22-24, W Africa, coll.

A. Murray. 1, BMNH 1891.4.2.14, Lagos, coll. Maloney. 3,

BMNH 1896.5.5.51-53, Lambarene, Ogowe R., coll. Kingsley. 1, BMNH 1900.6.28.311, St. Louis, Senegal, coll.

Delhez. 3, BMNH 1901.12.28.112.114, Gambia, coll. Budgett. 1, BMNH 1902.11.10.262, Lagama, S Nigeria, coll.

Ansorge. 1, BMNH 1911.6.1.131, Gunja, Angolia, coll.

Ansorge. 1, BMNH 1913.7.12.2, Ogowe, coll. Schneider. 1,

BMNH 1914.11.2.66, Lagos, coll. Cadman. 1, BMNH

1930.3.24.44, Accra, Gold Coast, coll. Irvine. 1, BMNH

1937.4.9.33, Nigeria, coll. Welman. 1, BMNH 1939.7.12.39,

Volta R., Gold Coast, coll. Irvine. 2, BMNH 1948.8.12.1-2,

Lagos, Lagoon, coll. Trewavas. 1, BMNH 1949.4.30.3, W

Africa, coll. Chabanaud. 1, BMNH 1953.7.10.41, Nigeria, coll.

Trewavas.

4. Cynoglossus dubius Day

FIGURE 15; PLATE 2

Cynoglossus dubius Day, 1873:525 [type-locality: Gwadur];

1877:435, pi. 95: fig. 2: 1889:456.—Norman, 1928:200.—

Saramma, 1963:75 [Kerala coast].—Pradhan, 1964:458 [Bombay coast].

DESCRIPTION.—Based on 8 specimens, 257.0-450.0

SL.

Depth of body 27.24-30.86 (M = 29.23), length of
NUMBER 238

km

FIGURE 15.—Outline drawing of C. dubius (BMNH 1911.12.6.16) from Karachi, Pakistan.

head 23.46-26.07 (M = 24.79) percent of standard

length. Diameter of eye 5.97-7.61 (M = 6.65), interorbital width pronounced, 7.46-9.72 (M = 8.42)

percent of length of head. Anterior nostril of eyed

side tubular, on upper lip in front of fixed eye,

the posterior in the anterior half of interorbital

space, or in the middle of it. Snout obtusely

pointed, 40.30-45.78 (M = 42.62) percent of length

of head, rostral hook rather short, extending to just

beyond mandibular symphysis. Maxillary extending to beyond posterior border of fixed eye; angle

of mouth extending below vertical from posterior

border of fixed eye, or a little beyond, nearer to

branchial opening than to tip of snout; snout to

angle of mouth 52.63-54.35 (M = 53.74), angle of

mouth to branchial opening 45.37-50.53 (M =

47.33) percent of length of head.

Scales: Cycloid on both sides, except along both

dorsal and anal fins toward the posterior side of

the ocular side, where the scales are weakly ctenoid.

Lateral-Line System: Two lateral lines on eyed

side, midlateral line with 98-104 scales, 17-21

scales between two lateral lines. One lateral line

on blind side. The midlateral line on ocular side

instead of opening through simple pores on every

scale opens by means of ducts into the adjoining

scale, generally one duct on one side followed by

three ducts on the other side (Figure 2).

Interlinear scale rows 17

Frequencies 3

18 19 20 21

S 11 0 1

Fins: Dorsal with 111-114 (M = 113) rays, anal

with 88-91 (90) rays, caudal 12 in 5 specimens

(radiographs).

Vertebrae: 52-53, comprising 9 abdominal and

43-44 caudal elements in 5 specimens (radiographs).

Coloration: Upper side uniformly brown, lower

whitish in preserved specimens.

Size: The largest specimen examined, 447.0

(450 + 27) mm, is from the Arabian Sea, trawled by

Anton Bruun (lat. 23°16/N, long. 67°56/E).

DISTRIBUTION.—West coast of India.

DIAGNOSIS AND AFFINITIES.—Cynoglossus dubius

is closely allied to C. borneensis. It is distinguishable, however, by having cycloid scales on the ocular side and by the greater number of anal fin rays

(88-91 cf. 82-87) and vertebrae (52-53 cf. 48-51).

NOTE ON SYNONYMY.—Originally described on the

basis of a single specimen by Day, C. dubius was

well diagnosed and there has been no confusion

with regard to its identity.

TYPE SPECIMEN.—Day (1873:525) described C.

dubius from a single specimen, 500 mm in TL, from

Gwadur and illustrated the specimen in his Fishes

of India (pi. 59: fig. 2). Norman (1928:200) described the species from four specimens, 220-460

mm in TL, and stated that it included an example

"believed to be the type of the species." Day's type

specimen was lost during the Varuna floods in

Banaras in 1943, where the collections of the Zoological Survey of India were removed for safety

during World War II.

OTHER MATERIAL EXAMINED.—1 specimen, ZSI (unregistered),

Kumta, Udipi, Mysore State, coll. Umarkutty and B. Sinha,

26.12.66. 1, USNM 139718, backwaters of Travancore. coll.

Schultz. 2, SOSC Ref 23, Neendakara, Kerala, lat. ca.

08a56'N, long. ca. 76°S0'E, coll. Anton Bruun, Oct. 1966. 4,

SOSC Ref 23. Arabian Sea. lat. 23'16'N, long. 67'56'E. coll.

Anton Bruun, 19.11.63. 1, BMNH 1906.10.24.18, Travancore,

ex. Trivandrum Museum. 1, BMNH 1911.12.6.16. Kurrachee

(Karachi), coll. Townsend. 1, BMNH 1912.7.20.28. Travancore, ex. Trivandrum Museum. 1, BMNH 1938.8.9.13. Calicut, 5 India, coll. Devanesan.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

5. Cynoglossus borneensis (Bleeker)

FIGURE 16; PLATE 3

Plagusia borneensis Bleeker, 1858a:6 [type-locality: Borneo].

Arelia borneensis.—Bleeker, 1860b:9.

Cynoglossus borneensis.—Bleeker, 1875:34, pi. 245: fig. 5.—

Gunther, 1862:498 [Singkawang].—Fowler, 1905:518.

[Borneo].—Seale, 1910:288.—Weber and de Beaufort,

1929:205.

Plagusia trulla Cantor, 1850:1213 [type-locality: Penang Sea,

Malay Peninsula].

Cynoglossus trulla.—Gunther, 1862:503.—Duncker, 1904:169.

Trulla cantori Kaup, 1858:109 [based on Cantor's P. trulla].

Cynoglossus sinicus Wu, 1932:146.

DESCRIPTION.—Based on 4 specimens, 88.0-300.0

ramSL.

Depth of body 23.04-25.50 (M = 24.28), length of

head 21.77-25.00 (M = 23.30) percent of standard

length. Diameter of eye 5.93-9.09 (M = 7.80), interorbital width 4.35-6.82 (M = 6.17) percent of length

of head. Anterior nostril of eyed side tubular, on

upper lip in front of lower eye, posterior nostril

km

FIGURE 16.—C. borneensis: a, outline drawing of lateral view of ocular side of specimen (BMNH

18j84.2-26.73) from China; b, details of anal region on blind side of specimen (BMNH

1931.4.23.53) from Borneo, female. (A = anus, OV = ovarian orifice.)
NUMBER 238

simple, in anterior half of interorbital space. Snout

rounded, 29.55-37.29 (M = 34.36) percent of length

of head, rostral hook short, extending hardly to

front of anterior nostril. Maxillary extending to beyond fixed eye; angle of mouth extending right up

to or just beyond vertical from posterior margin of

fixed eye, nearer to tip of snout than to branchial

opening; snout to angle of mouth 43.18-50.85

(M = 46.86), angle of mouth to branchial opening

51.49-54.55 (M = 53.44) percent of length of head.

Scales: Ctenoid on ocular side except those on

lateral lines, scales on blind side and those on

lateral lines of eyed side cycloid.

Lateral-Line System: Two lateral lines on eyed

side, midlateral line with 94-102 scales, 18-20

scales between the two lateral lines. One lateral

line on blind side.

Interlinear scale rows 18 19 20

Frequencies 1 2 1

Fins: Dorsal with 108-113 (M • 111) rays, anal

with 82-87 (M = 84) rays, caudal 12 in 8 specimens

(radiographs).

Vertebrae: 48-51, comprising 9 abdominal and

49-52 caudal elements in 8 specimens (radiographs).

Coloration: Upper side with three approximately

parallel longitudinal stripes, the middle one on

median lateral line and a diffuse dark patch on

opercular region, lower side whitish in preserved

specimens.

Size: Largest specimen examined, 450.0 mm SL,

is from China.

DISTRIBUTION.—Borneo, Malaya, Gulf of Thailand, and South China Sea.

DIAGNOSIS AND AFFINITIES.—Cynoglossus borneensis is closely related to C. dubius but can be readily

separated by the ctenoid scales on the ocular side

(cycloid in C. dubius) and the more elongate body

(depth of body 23.04-25.50 cf. 27.24-30.86 percent

ofSL).

NOTE ON SYNONYMY.—Bleeker (1858:6) described

P. borneensis on the basis of a single specimen, 218

mm TL from Sinkawang, Borneo, and characterized it as having two lateral lines on the ocular side

and a single one on the blind side. In the Atlas

(1875:34, pi. 245:fig. 5) he redescribed the species

and illustrated it in color. Because of his excellent

description and illustration there has not been any

confusion in the identity of C. borneensis. Cantor

(1850:1213) considered a specimen, 300 mm TL,

from the sea of Penang, Malay Peninsula, to represent a new species, Plagusia trulla, and characterized

it as having only one nostril in front of the lower

eye, the depth of the body four and one-half to

four and two-thirds in TL, and 12 rays in the

caudal fin. Cantor, however, did not mention the

number of lateral lines on the body. I have examined the type of P. trulla in the British Museum

and though the nostrils are not discernible in the

stuffed specimen smeared over with varnish, an

interlinear scale row of 18 or 19 scales and the 12

rays in the caudal fin are discernible, distinctive

enough to refer the specimen to C. borneensis.

Cynoglossus sinicus described from Chinese waters

is the same as the present species. Three specimens,

one from Hong Kong and the other two from

China labeled as C. sinicus in the British Museum,

agree so well in body depth and scales with C.

borneensis that I have no reservations in synonymizing them.

TYPE SPECIMENS.—Among the Bleeker collection

specimens in Leiden is a specimen, 195 mm SL, 212

mm TL, from Sinkawang, Borneo, catalogued as

RMNH 6792 (pers. comm., Dr. M. Boeseman); it is

evidently the holotype of C. borneensis.

There are two stuffed specimens, 212 and 262 mm

SL, of P. trulla cataloged as BMNH 1860.3.19.431

from Penang, coll. Dr. Cantor.

OTHER MATERIAL EXAMINED.—MALAY: 1 specimen, BMNH

1860.3.19.431a, Penang, Malay, coll. Dr. Cantor. 2, BMNH

1933.7.31.28-29, Singapore Mkt., ex. Raffles Museum.

BORNEO: 1, Kapuas R., Borneo, coll. Hardenberg. THAILAND: 1, BMNH 1862.11.1.225, Thailand, coll. Jamrach. 1,

USNM 103321, Bandon Blight, Gulf of Thailand, coll. H. M.

Smith, 29.9.1923. 1, CAS (GVF) 2582, Shakon Prov. y4 mi

off shore, Gulf of Thailand, lat. 13°26'15''N, long. 100° lOT.

coll. Naga Expd. 1959-1961. 7, ANSP 87208, Thailand, coll.

R. M. de Schauensee, 1936. CHINA: 1, BMNH 1839.353.39.

Hong Kong, coll. Herklots. 2. BMNH. 1884556.73, China.

The broumi complex

Cynoglossus browni is placed in a separate complex, which is close to the canariensis complex of

species but differs in the absence of a lateral line

on the blind side. Morphologically and in its distribution C. browni is related to C. senegalensis.

6. Cynoglossus browni Chabanaud

PLATES

Cynoglossus senegalensis morpha browni Chabanaud,

1949b:204 [type-locality: coast of Sierra Leone, West Africa].
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cynoglossus browni.—Cadenat, 1960:1382, pi. 2: fig. B; pi. 3:

figs, B, c.—Nijssen, 1966:87 [28 miles northwest of Ijmuiden,

off the coast of Netherlands].

DESCRIPTION.—Based on 12 specimens, 219.5-370.0

mm SL, including the holotype of C. browni.

Depth of body 24.22-28.53 (M = 25.93), length

of head 17.67-20.90 (19.02) percent of standard

length. Diameter of eye 3.74-6.60 (M = 4.81),

interorbital width rather broad 6.25-9.43 (M =

7.74) percent of length of head. Anterior nostril of

eyed side on upper lip in front of lower eye, posterior nostril in anterior half of interorbital space.

Snout narrowly rounded, 26.85-32.08 (M = 29.29)

percent of length of head, rostral hook short, not

reaching to vertical through front border of migratory eye. Maxillary extending well beyond fixed eye;

angle of mouth extending to beyond vertical from

posterior border of fixed eye and much nearer to tip

of snout than to branchial opening; snout to angle

of mouth 39.08-50.91 (M = 45.57), angle of mouth

to branchial opening 54.02-58.39 (M = 55.40) percent of length of head.

Scales: Ctenoid on ocular side except those on

lateral lines, scales on blind side and those on

lateral lines of ocular side cycloid.

Lateral-Line System: Two lateral lines on ocular

side, midlateral line with 84-91 scales, 14-16 scales

between middle and upper lateral lines. No lateral

line on blind side.

Interlinear scale rows 14 15 16

Frequencies 2 6 9

Fins: Dorsal with 119-125 (M = 121) rays, anal

with 96-99 (M = 97) rays, caudal 12 in 15 specimens

(radiographs).

Vertebrae: 57-59 comprising 9 abdominal and

48-50 caudal elements in 15 specimens (radiographs).

Coloration: Upper side more or less dark brown,

lower rather whitish in preserved specimens.

Size: The largest specimen examined, 402.0 (370

+ 32) mm, is from Nigeria, off Lagos.

DISTRIBUTION.—West Africa: Senegal to Nigeria

and Congo. The record from the Netherlands coast

(Nijssen, 1966) is a stray occurrence and the

northernmost limit is most likely Senegal.

DIAGNOSIS AND AFFINITIES.—Cynoglossus browni

shows affinities with C. senegalensis, C. canariensis,

and C. monodi but can be readily distinguished by

the total absence of a lateral line on the blind side.

TYPE SPECIMENS.—Holotype of C. senegalensis

browni, MNHP 49.23, 219.5 mm SL, from Sierra

Leone, west Africa.

OTHER MATERIAL EXAMINED.—10 specimens, USNM 188486,

off Lagos, Nigeria, coll. A. Longhurst. 6, USNM 188437, off

Nigeria, coll. A. Longhurst. 1, USNM 193914, off St. Paul R.,

Liberia, coll. George C. Miller. 1, ANSP 8690, Cape Palmas,

Liberia, coll. Thomas S. Savage.

The bilineatus complex

Cynoglossus bilineatus, the only species of the

bilineatus complex, is characterized by 12 rays in

the caudal fin and two lateral lines on the blind

side. Its most important difference from the members of the canariensis complex of species is an

additional lateral line on the blind side. The nature

of the scale pattern on the body is very similar

to that of the members of the canariensis complex,

especially in having cycloid lateral-line scales on the

ocular side, and in the interlinear scale formula of

13-17 scales. The distribution of C. bilineatus,

from the evolutionary center of the genus (Malay

Archipelago) westward to India and Pakistan, south

to New Guinea and Australia, and north to Taiwan

and Japan, suggests its probable origin in the Malay

Archipelago from an early stock of the canariensis

group.

7. Cynoglossus bilineatus (Lacepede)

FIGURE 17; PLATE 4

Achirus bilineatus Laclpede, 1802:6 [type-locality: China and

the East Indies].

Cynoglossus bilineatus.—Ogilby, 1910:39.—Weber, 1913b:443.

—Norman, 1926:301 [Queensland]; 1928:198.—Weber and

de Beaufort, 1929:194.—Herre, 1933:5 (Sandakan, N.

Borneo); 1934:105 [Manila].—Roxas and Martin, 1937:70

[Philadelphia].—Hardenberg, 1941:226 [Merauke, New

Guinea].—Suvatti, 1950:326.—Herre, 1953:189 [Philippines].—Munro, 1955:264, pi. 50: fig. 767; 1958:285

[Merauke].—Saramma, 1963:75 [Kerala coast].—Pradhan,

1964:458 [Bombay coast].—Marshall, 1964:468, pi. 64: fig.

454 [east coast of North Queensland].

Plagusia quadrilineata Bleeker, 1851a:412; 1852:21.

Arelia quadrilineata.—Kaup, 1858:107 [Java, Sumatra].—

Oshima, 1927:198 [Tainan, Taihoku].

Cynoglossus quadrilineatus.—Giinther, 1862:497.—Kner,

1867:295.—Bleeker, 1875:32, pi. 245: fig. 3.—Klunzinger,

1871:573.—Day, 1877 [in part]:435; 1880:409.—Madeay,

1884:53.—Alcock, 1889a:288.—Day, 1889 [in part]:457 —

Smith and Pope. 1906:498 [Kochi].—Jenkins, 1910a:30.—

Wu, 1932:144.
NUMBER 238 37

I cm

FIGURE 17.—Outline drawing ot C. btlineatus (BMNH 1931.4.23.52) from Java.

Cynoglossus lineolatus Steindachner, 1867:588 [Hong Kong].—

Bleeker, 1873:133.—Rutter, 1897:88 [Swatow].—Reeves,

1927:14.—Chu, 1931:95.—Wu, 1932:150.—Fowler. 1934b:218

[Hong Kong, Swatow].

Cynoglossus quinquelineatus Day, 1877:432, pi. 98: fig. 1

[type-locality: Madras]; 1889:453.—De Silva, 1956:198.—

Norman, 1928:197.

Cynoglossus sindensis Day, 1877:434, pi. 90: fig. 6 [typelocality: from Sind through the seas of India].—Jordan and

Richardson, 1908:281.—Ogilby. 1910:37 [Croker Island,

Northern Territory. Australia].—Norman, 1926:302.—De

Silva. 1956:198.

Arelia diplasios Jordan and Evermann, 1903:367, fig. 29

[type-locality: Formosa].—Jordan and Richardson, 1909:202,

fig. 25 [Formosa].—Oshima, 1927:204.

DESCRIPTION.—Based on 24 specimens, 95.0-338.0

mm SL, including the holotype of Cynoglossus

quinquelineatus Day.

Depth of body 22.22-28.60 (M = 25.86), length

of head 19.15-24.40 (M = 22.21) percent of standard

length. Diameter of eye 6.40-11.90 (M = 9.04),

interorbital width 5.0-10.71 (M = 7.86) percent of

length of head. Two nostrils on eyed side, anterior

one tubular, in front of lower eye. The posterior

nostril in anterior half or middle of interorbital

space. Snout rounded, 32.61-54.39 (M = 37.15),

rostral hook short, reaching hardly before vertical

through front border of anterior nostril. Maxillary

extending beyond fixed eye; angle of mouth extending to below vertical from posterior border of lower

eye, slightly nearer to branchial opening than to

tip of snout; snout to angle of mouth 46.44-56.90

(M = 51.38), angle of mouth to branchial opening

47.92-55.56 (M = 49.97) percent of length of head.

Scales: Ctenoid on ocular side except those on

lateral lines; scales of blind side and those of lateral

lines of ocular side cycloid.

Lateral-Line System: Two lateral lines on ocular

side,7 88-96 scales on median lateral line, 13-16

scales between the two lines. Two lateral lines on

blind side.

Interlinear scale rows

Frequencies

13 14 15 16

6 17 5 4

Fins: Dorsal with 107-113 (M = 110) rays; anal

with 80-88 (M = 86) rays, caudal 12 in 10 specimens

(radiographs).

Vertebrae: 51-53 comprising 9 abdominal and

42-44 caudal elements in 10 specimens (radiographs).

Coloration: Upper side in alcohol brownish with

an irregular dark patch on opercular region, lower

whitish.

Size; Largest specimen examined, 354.0 mm, is

from Colombo.

DISTRIBUTION.—From Malay Archipelago to the

coasts of India and Pakistan and to New Guinea

(Croker Island, Northern Territory) and coasts of

Queensland and Taiwan and Japan (Kuchi).

DIAGNOSIS AND AFFINITIES.—In the number of two

lateral lines on both sides of the body, C. bilineatus

approaches the attenuatus complex; it is immediately distinguished from these species by having 12

rays in the caudal fin, whereas the species of the

attenuatus complex have only 10. The interlinear

count of 13-16 in C. bilineatus (10-11 in C. attenuatus, 16-18 in C. lachneri, and 18-20 in C. dispar)

further distinguishes it from members of the

attenuatus complex.

'Day's C. quinquelineatus has an incomplete third lateral

line on the eyed side, a variation in the lateral-line system

not uncommon in the genus.
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

NOTE ON SYNONYMY.—Norman (1928:198) included C. quadrilineatus Bleeker and C. sindensis

Day in the synonymy of C. bilineatus. Ochiai (1963:

76) synonymized A. diplasios Jordan and Evermann

with C. bilineatus and I concur with them. Day

(1877:432) described C. quinquelineatus on the basis

of a single specimen in which an additional lower

lateral line on the ocular side is present. Day further

noted that his species differed from C. bilineatus

in having a somewhat smaller eye, a shorter rostral

hook, and in the position of the angle of the mouth,

which extends hardly to the posterior edge of the

eye. I have examined the holotype of C. quinquelineatus in Calcutta and found that, except in the

additional lower lateral line, it differs from C.

bilineatus in no other specific character. As has

been pointed out earlier, the nature and extent of

the lateral lines in Cynoglossus are of no significance

in specific differentiation, and hence the development of incomplete lateral line on the lower part of

the body in C. quinquelineatus is of no consequence

so I have synonymized it with C. bilineatus.

Cynoglossus lineolatus, described on the basis of a

single specimen, 106 mm in TL, from Swatow,

Hong Kong and characterized as having two lateral

lines on the blind side, is also referable to C.

bilineatus.

TYPE SPECIMENS.—The type of C. bilineatus is

not available in the Paris Museum (pers. comm.,

Madam Bauchot) and is not extant. Bleeker (1851a:

412) described Plagusia quadrilineatus on the basis

of 18 specimens, 75-300 mm in TL, from Batavia,

Sumatra, Muntok, and Bangka. In his Atlas (1875:

32, pi. 245: fig. 3), Bleeker described and illustrated

a specimen of 300 mm in TL. Among the Bleeker

collection of specimens in Leiden there are 9 specimens obtained from Batavia, west Sumatra, and

Boeling (Bali) and cataloged as RMNH 6789, none

of which matches either 75 or 300 mm. Since it is

difficult to ascertain which specimen should be

considered typical, all the 9 specimens in RMNH

are assumed to be syntypes of P. quadrilineatus

(Dr. M. Boeseman examined all the 9 specimens on

my behalf and recommended this solution).

NHV 43.781, 102 mm SL, 111 mm TL, from

Hong Kong, holotype of C. lineolatus.

Holotype of C. quinquelineatus, ZSI 1265, 234

mm SL, 250 mm TL, the figured specimen by Day

(1877, pi. 98: fig. 1).

OTHER MATERIAL EXAMINED.—JAPAN: 1 specimen, USNM

59754, Japan, coll. H. M. Smith. 1. USNM 59755. Japan,

coll. H. M. Smith. PHILIPPINES: 1. USNM 83396, Philippine Islands, coll. Hongkong Fisheries, 18.11.67. 1. USNM

113179. Limbones Cove, coll. Albatross. USNM 137652.

Manila Mkt., coll. Albatross. 3, USNM (unregistered), Limbones Cove, coll. Hongkong Fisheries Sta, 18.11.67. 1, MCZ

30567, Macasser, Celebes, coll. T. Barbour. 2, ANSP 49038-

39. Philippine. 1, NHV 43811. Manila, coll. Schilling, 1828.

1, BMNH 1872.1.18.18. Philippines, coll. Veitch. 7. USNM

137616-137620, Philippines and Borneo, coll. Albatross, 1907-

1909. BORNEO: 1, BMNH 1895.2.28.63, Sarawak, coll.

Brooke. INDONESIA: 3, USNM 72751. Java, coll. Bryant

Palmer. 16, USNM 159420, Java, coll. Hubbs and Hardenberg, 6.15-1929. 5, MCZ 30803, Batavia, Java, coll. Bryan

Palmer, 1909. 4, ANSP 772.74, Medan. Sumatra, coll. George

Vanderbilt Sumatra Expd., 23.5 1939. 1. BMNH 1858.4.29.3,

Amboyna, coll. Frank. 1, BMNH 1931.4.23.51. Batavia Mkt.,

coll. Hardenberg. 1, BMNH 1931.4.23.52. Java. coll. Hardenberg.

THAILAND: 2, ANSP 61829-31, Paknam, coll. R. M. de

Schauensee. 21.8.1934. 2, ANSP 61827-28, Bangkok, coll.

R. M. de Schauensee, 1734. 1, MCZ (unregistered), Thailand.

7

#

53'09"N. 98'50'07-E coll. BM Thai-Danish Expd., 15.2.1966.

1, NHV 43810, locality not given, coll. Hein. 1902. 1. CAS

(GVF). Gulf of Thailand, lat. 10°26'S4wN, long. 9°15'24"E,

coll. Thai Fishermen, Naga Expd., 1959-1961. 2. CAS (GVF),

Gulf of Thailand, coll. Thai Fishermen, Naga Expd., 24-

6.1960. 1, BMNH 1920.7.13.75, Thailand, coll. Prince Chumpen. SINGAPORE: 1, MCZ 32965, Singapore, coll. F. W.

Putnam, 1864. 2, BMNH 1933.7.31.33. 34, Singapore Mkt..

ex. Raffles Museum. AUSTRALIA: 1, BMNH 1925.7.22.82.

Queensland, coll. Endeavour. BURMA: 1. ZSI. 3444/1, Elephant point, Arakan coast, coll. Bengal Fisheries.

INDIA: 1, ZSI 12702. Ganjam coast, coll. Marine Survey.

2, ZSI 1145, 1146, Madras, coll. F. Day. 8, ZSI 1231-1234.

12263, Orissa coast, 7-8 fms, coll. Marine Survey. 1, ZSI

26227/1, Pun coast, Orissa. coll. Dr. A. Annandale. 2. ZSI

F 2360/2, Marina Beach, Madras, ex. Southern Regional Sta,

ZSI; Madras. 2, ZSI (unregistered), Karwar coast, Bombay,

coll. Karwar Survey 1967. 1, Maharastra. lat. 18M5'N, long.

72°25'E. coll. F. H. Berry. 1. MCZ 4273, Malabar, coll. F.

Day. 1, BMNH 1928.350.81, Madras Harbor, ex. Indian

Museum. 2, BMNH 1928.3.20.82, 83, Orissa coast, 7-8 fms,

ex. Indian Museum. 1, BMNH 1938.8.9.14, Calicut, coll.

Devanesan. PAKISTAN: 8. SOSC, W Pakistan, W end of

Astola Is., W of Karachi, lat. 25o06'35"N, long. 63°48'65"E,

coll. Anton Bruun (Woods, Fehlman, Rogers, Stone, and

Berry). 1, BMNH 1898.12.24.49, Karachi, coll. Townsend. 3.

BMNH 1911.12.6.14, Karachi, coll. Townsend. CEYLON: 1.

MCZ 33197, Colombo, coll. Capt. Putnam.

The attenuatus complex

Like C. bilineatus, the members of the attenuatus

complex are large species with two lateral lines on
NUMBER 238 39

the blind side but with only 10 rays in the caudal

fin. Cynoglossus attenuatus and its relatives C.

lachneri and C. dispar have their center of distribution in the central and western sectors of the

Indian Ocean and may have descended from the

same stock that gave rise to the bilineatus complex.

8. Cynoglossus attenuatus Gilchrist

FIGURE 18; PLATE 4

Cynoglossus attenuatus Gilchrist, 1905:11, pi. 29 [type-locality:

Tugela R. mouth, 4i/£ miles northwest, South Africa].

Arelia attenuata.—Gilchrist and Thompson, 1917:398.—Regan,

1920:221 [Natal].—Barnard, 1925:413 [Natal and Zululand

to Delagoa Bay].—Bonde, 1925:292.—Fowler, 1925:187;

1935:405.—Barnard, 1937:41.

Cynoglossoides attenuatus.—Bonde, 1922:23.

Arelia bilineata.—[not Lacepede], Smith, 1949 [in part]: 166,

pi. 2: fig. 341.

DESCRIPTION.—Based on 3 specimens, 158.0-244.0

mm SL, including the holotype.

Depth of body 21.20-25.59 (M = 23.36), length

of head 20.89-21.95 (M = 21.12) percent of standard

length. Diameter of eye 8.57-12.12 (M = 9.93),

interorbital width 4.55-6.06 (M = 5.44) percent of

length of head. Anterior nostril on eyed side in

front of lower eye, at a level with its lower margin,

posterior nostril in anterior half of interorbital

space. Snout obtusely pointed 36.36-43.94 (M =

40.10) percent of length of head, rostral hook reaching below anterior nostril. Maxillary extending

beyond fixed eye; angle of mouth extending to

below vertical from posterior border of fixed eye,

nearer to branchial opening than to tip of snout;

snout to angle of mouth 52.27-57.58 (M = 54.08),

angle of mouth branchial opening 47.73-50.48 (M

= 49.04) percent of length of head.

Scales: Ctenoid on ocular side, cycloid on blind

side; scales on lateral lines of ocular side ctenoid

anteriorly and cycloid posteriorly.

Lateral-Line System: Two lateral lines on ocular

side, median lateral line with 70-76 scales, 10-11

scales between them. Two lateral lines on blind

side with 11-12 scales between them.

Interlinear scale rows 10 11

Frequencies 1 1

Fins: Dorsal with 114-118 rays; anal with 90-91

rays, caudal 10 in 2 specimens (radiographs).

Vertebrae: 54-55, comprising 9 abdominal and

45-46 caudal elements in 2 specimens (radiographs).

Coloration: Upper side uniformly brown, lower

whitish in preserved specimens.

Size: The largest specimen examined, 310 (290 +

10) mm, is from Durban, South Africa.

DISTRIBUTION.—South Africa: Natal and Zululand

to Delagoa Bay.

DIAGNOSIS AND AFFINITIES.—Cynoglossus attenuatus, which is restricted in its distribution to South

Africa, is related to C. lachneri and C. dispar.

Phylogenetically, it is most probable that C. attenuatus was derived from a population of the early

generalized stock of the bilineatus-attenuatus complex in South African waters. The attenuated nature

of body (mean depth of 23 percent of SL, cf. 25.95

of C. lachneri and 29.87 of C. dispar) and large

scales (10-11) distinguish it from the other species

(16-18 in C. lachneri and 18-20 in C. dispar).

TYPE SPECIMEN.—BMNH 1903.12.31.10, holotype

of C. attenuatus, 210 mm SL, 4i/2 miles, NW of

south head of R. Tugela, Natal, 24 fans, coll.

Gilchrist.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH

I cm

FIGURE 18.—Outline drawing of C. attenuatus (BMNH 1919.9.1231) from Durban.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1919.9.12.51, Durban, coll. Marley. 2, ANSP 88867, Delagoa

Bay. South Africa, coll. H. W. Bell Harley, 1923.

9. Cynoglossus lachneri, new species

PLATE 5

DESCRIPTION.—Based on 12 specimens (130.0-4

32.0 mm SL), including the holotype and paratypes.

Depth of body 24.62-27.19 (M = 25.95), length

of head 17.48-23.14 (M = 19.43) percent of standard

length. Diameter of eye 7.41-10.89 (M = 9.34),

interorbital width 6.98-9.76 (M = 8.20) percent of

length of head. Anterior nostril of the eyed side

tubular, on the upper lip, situated almost vertical

through the anterior border of upper eye, and some

distance in front of lower eye, posterior nostril a

simple opening, in anterior half of interorbital

space. Two nostrils on blind side, the tubular

anterior one on the anterior half of the upper lip,

the posterior a little higher and above posterior

half of upper lip. Snout rounded, 28.40-34.26 (M =

31.47) percent of length of head; rostral hook

rather short, scarcely reaching vertical through front

border of migratory eye. Maxillary extending

beyond hind border of fixed eye; angle of mouth

extending below vertical from hind border of fixed

eye, nearer to tip of snout than to branchial opening; snout to angle of mouth 45.73-50.00 (M =

47.02), angle of mouth to branchial opening 50.00-

55.81 (M = 52.84) percent of length of head.

Scales: Ctenoid on ocular side, except those on

lateral lines; scales on blind side and those on

lateral lines of ocular side cycloid.

Lateral-Line System: Two lateral lines on eyed

side, midlateral line with 100-111 scales, 16-18

scales between the two lines. Two lateral lines on

blind side.

Interlinear scale rows 16 17 18

Frequencies 3 7 2

Fins: Dorsal with 113-121 (M = 117), anal with

92-98 (M = 96) rays, caudal 10 in 7 specimens

(radiographs).

Vertebrae: 55-58, comprising 9 abdominal and

46-49 caudal elements in 7 specimens (radiographs).

Coloration: Upper side uniformly dark brown,

lower whitish in preserved specimens.

Size: Largest specimen examined, 461.0 (432 +

29) mm, is from Mombasa, the holotype of lachneri.

DISTRIBUTION.—Mozambique (Inhaca) coast northward to Red Sea and Gulf of Oman and eastward

to Anjouan (Comoro) and Nossi-Be Islands and

Seychelles.

DIAGNOSIS AND AFFINITIES.—Cynoglossus lachneri

has close affinities with C. attenuatus and C. dispar

and in many characters is intermediate between

these two species. It is distinguishable, however, by

its interlinear scale count (16-18 cf 10-11 in C.

attenuatus, and 18-20 in C. dispar).

TYPE SPECIMENS.—USNM 201852. holotype of C.

lachneri, 432 mm SL, Mombasa, Kenya, coll. Anton

Bruun. 16.11.1964. Paratypes: 1 specimen, USNM

203963, paratype of C. lachneri, 285 mm SL, Sandy

Cove behind reef of Nosi N' Tanga Nossi-Be,

Madagascar, lat. 13°22'07"S, long. 48°10/45//E,

coll. Anton Bruun. 1, USNM 171046, paratype

of C. lachneri, 270 mm SL, Madagascar, coll. Dr.

J. Millot, Feb 1953. 1, MCZ 11551, paratype of

lachneri, 234.5 mm SL, Zanzibar, coll. Cooke. 1,

NHV 43874, paratype of C. lachneri, 130 mm SL,

Red Sea, coll. Staind; 1895-1896. 1, RU, paratype

of C. lachneri, 190 mm SL, Tiwi, Gulf of Oman,

coll. Mrs. J. L. B. Smith. 1, RU, paratype of C.

lachneri, 226 mm SL, Silhoutla Bay, Seychelles, coll.

J. L. B. Smith. 3, ZSI F 6 i2i , paratype of C.

lachneri, 176-274 mm SL, Pinda Beach, Mozambique, coll. J. L. B. Smith. 1, ZSI 620 7

 , paratype of

C. lachneri, 326 mm SL, Red Sea, Harkiko Bay, coll.

H. Steinitz. 1, ZSI F §|°-*L, paratype of C. lachneri,

227 mm SL, Inhaca, Mozambique, coll. Mrs. J. L. B.

Smith, Aug 1948.

OTHER MATERIAL EXAMINED.—2 specimens, BMNH

1869.3.1.30-31, Seychelles, coll. Wright. 1, BMNH

1932.73.53, Seychelles, coll. Wood.

10. Cynoglossus dispar Day

FIGURE 19; PLATE 5

Cynoglossus dispar Day, 1877:434, pi. 96: fig. 2 [type-locality:

Bombay]; 1889:456.—Norman, 1928:199 [Bombay, Madras].—Pradhan, 1964:458 [Bombay coast].

DESCRIPTION.—Based on 12 specimens, 167.0-350.0

mm SL, including the lectotype and paralectotypes

of C. dispar.

Depth of body 24.81-32.58 (M = 29.87), length

of head 18.83-21.35 (M = 20.17) percent of standard

length. Diameter of eye 7.81-9.91 (M = 8.75), inter-
NUMBER 238 41

FIGURE 19.—Outline drawing of C. dispar (BMNH 1889.2.1.4062) from Madras.

orbital width 4.48-9.01 (M = 5.95) percent of length

of head. Two nostrils on ocular side, a simple one

in anterior half of interorbital space, and a tubular

one in front of lower eye. Snout rounded 16.42-

31.53 (M = 26.60) percent of length of head, rostral

hook short and not extending to vertical through

the front border of upper eye. Maxillary extending

to beyond posterior border of fixed eye; angle of

mouth extending to about or almost below vertical

from hind border of fixed eye, much nearer to tip of

snout than to branchial opening; snout to angle of

mouth 38.81-52.25 (M = 43.50), angle of mouth

to branchial opening 46.85-58.73 (M = 56.0) percent of length of head.

Scales: Ctenoid on ocular side except those on

lateral lines; scales of blind side and those of

lateral lines of ocular side cycloid.

Lateral-Line System: Two lateral lines on eyed

side, 102-119 scales on median lateral line, 18-20

scales between them. Two lateral lines on blind

side.

Interlinear scale rows

Frequencies

18 19 20

2 3 S

Fins: Dorsal with 109-113 rays, anal with 90-92

rays, caudal 10 in 5 specimens (radiographs).

Vertebrae: 53-54, comprising 9 abdominal and

44-45 caudal elements in 5 specimens (radiographs).

Coloration: Upper side brownish with somewhat

darker irregular blotches, lower whitish in preserved

specimens.

Size: Largest specimen examined, 389 (350 + 33)

mm, is from Karachi, west end of Astola Island,

trawled by Anton Bruun.

DISTRIBUTION.—India: Madras and Bombay;

Pakistan: Astola Island, west of Karachi.

DIAGNOSIS AND AFFINITIES.—Cynoglossus dispar

is closely allied to C. lachneri but it can easily be

separated by its deeper body (mean depth of body

29.87 percent of SL cf. 25.95 percent of SL) and

larger scales (18-20 cf. 10-11 in C. attenuatus and

16-18 in C. lachneri).

NOTE ON SYNONYMY: Originally described by

Day from specimens from Bombay and Madras, C.

dispar was well diagnosed and there has been no

confusion with regard to its identity.

TYPE SPECIMEN: Day (1877:434) described C.

dispar from an unstated number of specimens from

Bombay and Madras. Norman (1928:199) redescribed the species from 4 specimens, 95-350 mm in

TL, including examples believed to be the types of

the species. Of the register numbers ZSI 1141 and

ZSI 1144, which Norman had listed as seen, only

the specimen bearing number ZSI 1141 is at present

available in the collections of the Zoological Survey

of India. The other one was presumably lost in

the Varuna floods of Banaras in 1943, where the

collections were temporarily shifted during World

War II. There are at present six specimens of the

syntype series that are traceable; including the one

lost, there must have been seven in the series. A

specimen, ZSI 2715, 260 mm SL, from Sind collected

by F. Day, which formed the basis for Day's figure

(pi. 96: fig. 2), is selected here as the lectotype, and

the rest of the five specimens ZSI 1141, Bombay,

coll. Day, BMNH 1889.2.1.4061, Sind, coll. F. Day,

BMNH 1889.2.1.4062, Madras, coll. F. Day, BMNH

1889.2.4063, Bombay, coll. F. Day, NHV 482 Madras,

coll. F. Day are selected as paralectotypes.

OTHER MATERIAL EXAMINED.—INDIA: 11 specimens, BMNH

1928.3.20.84, Bombay, ex. Indian Museum. 1. BMNH

1933.1.2.6, S. Malabar, coll. Devanesan. PAKISTAN: 5.
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SOSC Ace 23, Karachi, west end of Astola Is., lat. 25°06'35"N,

long. 63°48'65"E.

The kopsi group

Several related species complexes of Cynoglossus

may be assembled together in the kopsi group,

kopsi being closest to the evolutionary stem of this

group. They are typified by the smaller adult size,

large eyes, either contiguous or with a very narrow

interoribital space, two nostrils, the posterior one

occasionally hidden under scales or absent (sealarki

and itinus complexes), short obtusely pointed snout

with the angle of the mouth situated nearer to tip

of snout than to branchial opening, one to three

lateral lines on ocular side, the dorsolateral and

the ventrolateral either absent or incomplete, no

lateral line on blind side, 10 or 8 rays in the caudal

fin, and large scales. The absence of a lateral line

on the blind side, the reduced number of rays in

the caudal fin, the contiguous or subcontiguous

nature of the eyes, and the absence of the posterior

nostril in the sealarki and itinus complexes are

considered as specializations.

Included in this group are the kopsi, itinus,

ogilbyi, ecaudatus, and sealarki species complexes.

Their combined range extends to almost the entire

distributional limits of the genus.

The kopsi complex

This complex includes the morphologically most

generalized species of the group. These species have

7 to 12 interlinear scale rows, both the anterior or

posterior nostrils present, and the dorsolateral line

extending to varying distances along the dorsal

contour (when the ventrolateral line is absent) or

with an uninterrupted dorsolateral line (when the

ventrolateral is present).

Included in this assemblage are C. kopsi, C.

interrnptus, and C. joyneri. Its center of distribution covers the whole of the Indo-Australian Archipelago, northward through the Philippines to

Taiwan and Japan, and westward through the seas

of India and Pakistan to Cargados Garajos, the

Seychelles, and the Persian Gulf.

11. Cynoglossus kopsi (Bleeker)

FIGURE 20; PLATE 6

Plagusia kopsii Bleeker, 1851b:494 [type-locality: Rio in

Bintang Island, southeast of Singapore]; 1852:31; 1854a:99.

Arelia kopsi.—Bleeker, 1859:184.

Cynoglossus kopsii.—Giinther, 1862:493 [East Indian Archipelago].—Bleeker, 1875:31. pi. 241; fig. 3—Weber and de

Beaufort, 1929:189.

Cynoglossus kopsi morpha diagramata Chabanaud [in part],

1951d:273 [type-locality: Arafura Sea].

Cynoglossus brachycephalus Bleeker, 1875:38, pi. 244; fig. 6

[type-locality :Sumarta].—Regan, 1908:235 [Maldives and

Cargados Carajos].—Weber, 1913b:441.—Norman [in part],

1928:211.—Weber and de Beaufort, 1929:207 [W. Sumatra].—Munro, 1955:265, fig. 772 [Pearl Banks].—De Silva,

1956:199 [Aripu].—Fowler, 1956:188.—Saramma. 1963:78

[Kerala Coast].—Chen and Weng, 1965:93, fig. 64 [Tungkong].

Brachycephalus {Cynoglossus) brachycephalus.—Chabanaud,

1951b:78.

Cynoglossus praecisus Alcock, 1890b:442 [type-locality: off

Ganjam coast]

Cynoglossus breviroslris [not Day].—Johnstone, 1904:209;

Chen and Weng, 1965:92, fig. 63 [Tainan].

Cynoglossus venicolor Alcock, 1890b:442 [type-locality: Orissa

coast].—Alcock; 1896:330.—Norman, 1928:195.—Punpoka,

1964:74 [Chumpora Province, Thailand].

Cynoglossus sibogae Weber, 1913b:442.—Weber and de

Beaufort, 1929:200, fig. 54.—Herre, 1934:105 [Banang Sur,

La Union Province, Luzon]; 1953:191.—Punpoka, 1964:71

[Goh Korma].

Cynoglossus interruptus [not Giinther].—Chen and Weng,

1965:90, fig. 61 [Pescadores, Taiwan].

Cynoglossus melanopterus [not Bleeker] Shen, 1929:21 [typelocality: Hong Kong].

DESCRIPTION.—Based on 43 specimens, 50.0-177.0

mm SL, including the paralectotype of Weber's C.

sibogae, the holotype of C. versicolor, and the holotype of C. kopsi diagramata.

Depth of body 22.43-33.87 (M = 26.60), length

of head 15.84-24.59 (M = 21.47) percent of standard

length. Diameter of eye 11.76-19.05 (M = 14.22)

percent of length of head, interorbital space absent.

Two nostrils on ocular side, the anterior tubular in

front of lower eye, the posterior nostril simple and

in front of the interorbital space, sometimes hidden

under scales. Snout rounded 24.24-40.62 (M =

28.80) percent of length of head; rostral hook short,

extending to front of anterior nostril. Maxillary

extending to below posterior half of fixed eye;

angle of mouth extending to below vertical from

anterior half of fixed eye, much nearer to tip of

snout than to branchial opening; tip of snout to

angle of mouth 35.42-59.38 (M = 41.83), angle of

mouth to branchial opening 48.48-68.75 (M =

57.80) percent of length of head.

Scales: Ctenoid on both sides, including those of

lateral lines.
NUMBER 238 43

FIGURE 20.—Outline drawing of C. kopsi showing variation in number and extent of lateral

lines: a, specimen 67.5 mm SL (BMNH 1934.9.6.7) from Singlap, Singapore; b, specimen 11.32

mm SL (BMNH 1890.2.26.147) from Arafura Sea; c, specimen 150.0 mm SL (BMNH 1933.7.31.30)

from Singapore.

Lateral-Line System: Usually two or three lateral

lines on ocular side, dorsolateral line undulating

and extending along the dorsal contour of body

to varying distances, midlateral line with 57-72

(M = 60) scales, 7-12 (M = 9) scales between the

upper and the middle lateral lines. Ventrolateral

line sometimes present, extending to varying distance. No lateral line on blind side.

Interlinear scale rows 7 8 9 10 11 12

Frequencies 4 5 14 4 2 S

Fins: Dorsal with 103-115 (M = 107) rays, anal

with 80-91 (M = 82) rays, caudal 10 in 30 specimens

(radiographs).

Vertebrae: 50-55, comprising 9 abdominal and

41-46 caudal elements in 30 specimens (radiographs).

Coloration: Upper side brownish, generally with

irregular darker spots, lower whitish in preserved

specimens.

Size: Largest specimen examined, 187 (177 + 10)

mm, from China Sea near Hongkong, Albatross

collection.

DISTRIBUTION.—From Indo-Australian Archipel-
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ago to Philippines and Taiwan and westward to

seas of India and Persian Gulf.

DIAGNOSIS AND AFFINITIES.—Cynoglossus kopsi and

C. interruptus are allopatric species resembling each

other in their relatively small size, relatively large

contiguous eyes, short and somewhat obtusely

pointed snout, two or three interrupted lateral lines

on ocular side, and 10 rays in the caudal fin; C.

kopsi is distinguishable, however, by its larger scales,

the interlinear scale count being 7-12, M = 9 cf.

11-12, M = 11 in C. interruptus and by usually

having two lateral lines, the ventrolateral line occurring only rarely.

NOTE ON SYNONYMY.—Bleeker (1851b:494) described C. kopsi from Rio in Bintang, off southeastern Singapore and characterized it as having

subcontiguous eyes, three lateral lines on the ocular

side, and none on the blind side, without indicating,

however, the number of scale rows between the

middle and the upper lateral lines. Because of the

omission of the key character of interlinear scale

row from the original description of C. kopsi, there

has been considerable confusion in the identity of

it. In his Atlas (1875, pi. 241: fig. 3) Bleeker, however, gave an excellent illustration and indicated

nine rows of interlinear scale rows. In his description of C. brachycephalus (1875:38) from Sumatra,

Bleeker again delineated this species with subcontiguous eyes and two nostrils but with only two

later lines on the ocular side. In this case also,

Bleeker made no mention of the number of interlinear scale rows present in the species, though in

his illustration (1875, pi. 244: fig. 6) nine rows can

be distinctly seen. Thus, the presence of an additional lateral line in C. kopsi is the only difference

distinguishing it from C. brachycephalus. An additional lower lateral line as has been stated earlier

is of no importance in species differentiation in

Cynoglossus and I have, therefore, considered these

two species as synonymous.

Norman (1928:211) included C. praecisus and C.

sibogae in C. brachycephalus and concluded that

"this species is very closely related to C. kopsi

Bleeker, with which it may prove to be identical.

There is no trace, however, of a lower lateral line

on the ocular side in any of the specimens described

above." I have examined the types of C. praecisus

and C. sibogae and I corroborate Norman's tentative conclusion that all three of these species, C.

praecisus, C. sibogae, and C. brachycephalus, are

synonymous with C. kopsi.

Alcock (1890b:442) described C. versicolor, based

on a single specimen obtained by RIMS Investigator

off the Orissa coast, and characterized it as having

a single nostril and 12 series of scales between the

upper and middle lateral lines. On a reexamination

of the type, Norman (1928:195) noted the presence

of posterior nostril and traces of an incomplete

lower lateral line, characters distinctive of C. kopsi.

I have examined the holotype of C. versicolor and I

consider it here as conspecific with C. kopsi.

Cynoglossus melanopterus Shen described on the

basis of eight specimens, 120 to 152 mm in TL,

from Talo Harbour, Hongkong is the same as the

present species. From the description of the fish as

having small subcontiguous eyes, two nostrils, three

lateral lines on ocular side, and ctenoid scales on

the body, it does not appear to differ a bit form

C. kopsi and I have, therefore, synonymized them.

TYPE SPECIMENS.—Bleeker's original description

of C. kopsi was based on two specimens, 125 and

130 mm in TL, and he illustrated the 125-mm

specimen in the Atlas. In the Bleeker collection of

specimens there is only one specimen, though three

are recorded in the register and in the card index

and two in the auction catalog of 1879 (pers.

comm., Dr. M. Boeseman). A specimen 119 mm SL,

125 mm in TL, cataloged as RMNH 7680. Rio

(Riau Archipelago), coll. G. F. de Bruyn Kops,

1851, is selected here as the lectotype of C. kopsi.

Cynoglossus brachycephalus was based on two

specimens, 105 and 125 mm in TL, and the illustration drawn from the 125-mm specimen. Chabanaud

(1946) selected a specimen, 118 mm SL approximately 124 mm TL, from west Sumatra, RMNH

17877, as lectotype of C. lectotype of C. brachycephalus; it was taken from a jar of Bleeker's

collection from the 1879 auction cataloged as SMNH

6800 containing five specimens. Weber (1913b:442)

described C. sibogae on the basis of two specimens,

77 and 61 mm, from Molo Strasse (Siboga Sta 51)

and Lirung, Insel Saliban (Siboga Sta 133), respectively. The Molo Strasse specimen, 77 mm, which

is figured, is selected here as the lectotype and the

other specimen the paralectotype. Both the types

are in the Zoological Museum, Amsterdam (pers.

comm., Dr. N. Nijssen), but I have examined only

the paralectotype. Holotype of C. praecisus Alcock,

ZSI 12843, 111 mm SL, from Ganjam coast, coll.
NUMBER 238 45

Marine Survey. Holotype of C. versicolor, ZSI 12895,

116 mm SL, from Orissa coast, coll. Marine Survey.

Holotype of C. kopsi diagramata, BMNH 1890.2.-

26.147, 109 mm SL, from Arafura sea, coll.

Challenger.

OTHER MATERIAL EXAMINED.—CHINA: 1 specimen, USNM

113181, China Sea near Hongkong, coll. Albatross. 1, USNM

137982, China Sea, near Hongkong, coll. Albatross. 3, USNM

192859, Ma Kung Mkt., Taiwan, coll. Kuntz and Wells. 2,

SUSC Ref 340, off Teluk Kau, Halmahera, lat. 01°08.6'N,

long. 128°01'E, 30-25 fms coll. Tevega Expd. 2, SOSC Ref

340, South China Sea, off Sarawak, lat. 4°44'N, long. 113°23'E,

100 mm coll. Tevega Expd. 1, BMNH 1855.9.19.1247, China,

coll. Belcher. 1, BMNH 1862.6.3.18, China, coll. P. Bleeker.

1, BMNH 1935.4,18.7, South China Sea, coll. Lin.

PHILIPPINES: 11, USNM 112872 Illoilo, coll. Kuntz and

Wells. 1, USNM 112873, Illoilo, coll. Kuntz and Wells. 3,

USNM 112874, Illoilo, coll. Kuntz and Wells. 1, USNM

113178, Antonis Is., coll. M. Weber, Aug 1899. 1, USNM

137653, Towi, coll. Albatross. 1. USNM 137691, Ragay R.,

Ragay Gulf, coll. Albatross. 74, USNM 148586, Illoilo, coll.

Keller. SINGAPORE: 1. BMNH 1933.7.31.30, Singapore, ex.

Raffles Museum.

INDONESIA: 1, BMNH 1879.5.14.81, Arafura Sea. coll.

Challenger. 2, UZMK (unregistered), Java Sea, lat. 5°08'S,

long. 106°22'E, 35 m, coll. Th. Mortensen, 26.7.22. 1, UZMK

(unregistered), Java, lat. 5°53'S, long. 05°34'E, 25 m, coll.

Th. Mortensen. 1, UZMK (unregistered), Java Sea, 12 m,

coll. Th. Mortensen, 5-8-22. 1, UZMK (unregistered), Java

Sea, lat. 106°03'E, long. 5°23'S, coll. Th. Mortensen, 6.8.22.

1, UZMK (unregistered), Java Sea, lat. 5°57'S, long. 106°84'E,

22 m, coll. Th. Mortensen, 7.8.1922. 1, UZMK (unregistered).

Amboina, ca. 50 fms, coll. Th. Mortensen, 2.11.11. 1, UZMK

(unregistered), Amboina, coll. Th. Mortensen, 24.11.22. 1.

UZMK (unregistered), Java Sea, lat. 8°26'S, long. 114°29'E,

70 m, coll. Th. Mortensen, Java-S Africa Expd., 5.4.1929.

THAILAND: 1, CAS (GVF) 2312, Chumphon Prov. Gulf

of Thailand, coll. Naga Expd. 1959-1961. 10, CAS (GVF)

2312, Gulf of Thailand, lat. 12O16'45"N, long. 100°07'15"E,

coll. Naga Expd., 1959-1961. INDIA: 1, SOSC Ref 381,

Pondicherry, Madras State, coll. F. H. Berry, 5.10.66. 4,

BMNH 1928.3.20.101-104, Madras coast, 20 fms, ex. Indian

Museum. 1, BMNH 1928.3.20.106, Ganjam coast, ex. Indian

Museum. 10, ZSI 13021-34 Madras coast, 20 fms, coll.

Marine Survey. PERSIAN GULF: 1, ZSI (unregistered), Persian Gulf, 20°20'N, 48°47'E, 19.11.190, Marine Survey Sta

352, coll. Investigator. 1, UZMK (unregistered), Persian

Gulf, ca. 50 fms, 1937-1938. MADAGASCAR: 2, SOSC Ref

170, lat. 16°IPS, long. 43°53'E, 62 m, coll. Anton Brunn,

18.10.64. 1, SOSC Ref 170, lat. 16°03'S, long. 44°09'E, coll.

Anton Brunn. MALDIVE ISLANDS: 1. BMNH 1901.1231,

120, Maldives, coll. Gardiner. ST BRANDON GROUP: 4,

BMNH 1908.3.23.149-52, Cargados Garajos, coll. Gardiner.

MERGUI ARCHIPELAGO: 1, ZSI (unregistered), Mergui

Archipelago 12°35'N, 38°I6'E, Marine Survey Sta 552, coll.

Investigator.

12. Cynoglossus interruptus Giinther

FIGURE 21

Cynoglossus interruptus Giinther, 1880:70, pi. 30: fig. B

[type-locality: Yokohama, Japan].—Steindachner, 1896:220

[Kobe, Hioga, Nagasaki].—Jordan and Snyder, 1900:380

[Tokyo]; 1901:123 [Yokohama].—Smith and Pope, 1906:

498.—Wu, 1932:146.—Ochiai, 1959:202; 1963:82, pi. 17:

fig. 2.—Chu, 1963:538, fig. 403.—Shen, 1967:212 [Hong

Kong].—Ochiai, 1966:84.

Areliscus interruptus.—Jordan and Starks, 1906a:240 [Tokyo,

Nagasaki, Wakanoura, Matsushima, Onomichi].—Snyder,

1912:441.—Jordan, Tanaka, and Snyder, 1913:336.—

Jordan and Thompson, 1914:213.—Hubbs, 1915:494.—

Jordan and Hubbs, 1925:302 [Toba, Kobe, Mikawa.

Misaki].—Reeves, 1927:14.—Ui, 1929:276.—Schmidt and

Lindberg, 1930:1149.—Chu, 1931:95.—Kamohara, 1934:

462:—Kuronuma, 1939:83.—Yanai, 1950:22.—Kuroda, 1951:

390.—Honma, 1952:225.—Mori, 1952:184.

Cynoglossus nigropinnatus Ochiai, 1959:205 [type-locality:

Owase, Mimase, Kochi, Japan]; 1963:85, pi. 19.

FIGURE 21.—Outline drawing of C. interruptus, holotype (BMNH 1879.5.14.92)

from Yokohama, Japan.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DESCRIPTION.—Based on 34 specimens, 89.0-158.0

nun SL.

Depth of body 25.47-35.62 (M = 28.97), length

of head 18.52-24.11 (M = 21.25) percent of standard

length. Diameter of eye 10.53-21.05 (M = 14.18)

percent of length of head, interorbital space absent.

Two nostrils on ocular side, anterior nostril tubular

in front of fixed eye, posterior nostril simple, just

above anterior border of lower eye. Snout rounded,

25.0-37.14 (M = 31.81) percent of length of head,

rostral hook short, extending only to front of anterior nostril. Maxillary extending to just below middle of fixed eye; angle of mouth extending to below

vertical from anterior half of lower eye, much nearer

to tip of snout than to branchial opening; tip of

snout to angle of mouth 33.33-52.63 (M = 43.84),

angle of mouth to branchial opening 36.36-63.16

(M = 56.02) percent of length of head.

Scales: Ctenoid on both sides, including those on

lateral lines; scales small, moderately serrated with

short ctenii on blind side, weakly serrated on ocular

side.

Lateral-Line System: Two or three lateral lines

on ocular side. Dorsolateral line undulated, interrupted at middle of body, becoming obscure or

runnng backward along the dorsal contour entering

dorsal fin along 8th to 12th rays counted from the

rear; the ventrolateral line absent or present, when

present either interrupted at middle of body or

running backward and entering anal fin a few

rays counted from the rear. The dorsolateral line

is usually well developed when the ventrolateral

line is present to the fullest extent. Midlateral line

with 60-67 (M = 63) scales, 11-12 scales between

middle and upper lateral lines. No lateral line on

blind side.

Interlinear scale rows 11 12

Frequencies 17 14

Fins: Dorsal with 103:110 (M = 107) rays, anal

with 84-86 (M = 85) rays, caudal 10 in 5 specimens

(radiograph).

Vertebrae: 52, comprising 9 abdominal and 43

caudal elements in 5 specimens (radiograph).

Coloration: Upper side brownish, with or without

dark blotches, blind side yellowish in preserved

specimens.

Size: Largest specimen examined, 168 (158 +

10) mm, is from Mimase, Kochi, Japan.

DISTRIBUTION.—Japan.

DIAGNOSIS AND AFFINITIES.—Cynoglossus interruptus, which is restricted to Japan, can be readily

separated from C. kopsi by its smaller scales (11-12,

M = 11 cf 7-12, M = 9) and usually by the occurrence of a ventrolateral line which is absent in the

holotype.

NOTE ON SYNONYMY.—Gunther (1880:70) described C. interruptus on the basis of two specimens,

the largest being 150 mm in TL and characterized

it as having two lateral lines, upper reaching only

two-thirds to caudal, and 12 rows of scales between

the middle and the upper. A similar species, C.

nigropinnatus, was described by Ochiai on the basis

of 84 specimens, 80.5-211 mm SL, from Owase,

Kochi, and Mimase and was characterized by having

three lateral lines on the ocular side, the middle and

the upper being separated by 10 rows of scales, and

with ctenoid scales on the body. Comparison of

numerous specimens of C. interruptus has shown

beyond doubt that the depth of the body and the

presence of an additional lower lateral line noted

by Ochiai fall within the normal range of variation

of C. interruptus. I have, therefore, included C.

nigropinnatus in the synonymy of C. interruptus.

TYPE SPECIMENS.—Holotype of C. interruptus,

BMNH 1897.5.14.92, 140 mm SL, from Yokohama,

Japan, coll. Challenger; paratype, BMNH 1890.7.

26.146, 140 mm SL, obtained along with the holotype. One paratype of C. nigropinnatus, KU 18336,

158 mm SL, two paratypes KU 36619 and KU 26620,

156 and 131 mm SL, Mimase, Japan, coll. Kyoto

University.

OTHER MATERIAL EXAMINED.—CHINA: 1 specimen, BMNH

1848.3.16.195, China, coll. Belcher. JAPAN: 2, BMNH

1902.10.31.146-47, Japan, coll. Gordon Smith. 10, BMNH

1923.2.26.650-59, Tokyo, coll. Jordon. 8, BMNH 1923.2.26.-

660-667, Wahanoura, coll. Jordon. 30, USNM 56360,

Nagasaki, col. Bureau Com. Fisheries. 1, USNM 59757,

Kagoshima, coll. H. M. Smith. 1, USNM 59756. Kochi, colL

H. M. Smith. 1, USNM 71955, Suruga, coll. Albatross. 1,

USNM 73868, Yenoshima. 1, USNM 77065, Japan, coll.

Albatross. 8, USNM 151810 Kobe, coll. D. S. Jordan and

Yamamoto, 1922. 5, USNM 152480, Kobe, coll. D. S. Jordon,

1922. 1, NHV 43777, Tokyo, coll. Doderlein.

13. Cynoglossus joyneri Gunther

FIGURE 22; PLATE 6

Cynoglossus joyneri Gunther, 1878:486 [type-locality: Tokei,

Japan]; 1880:70 pi. 30; fig. A [Tokyo].—Otaki, 1897:25.—

Wu, 1932:154; 1933:61 [Chefoo].—Wu and Wang, 1933:304

[China].—Chabanaud, 1951d:269.—Tchang, 1955:301, fig.
NUMBER 238 47

FIGURE 22.—Outline drawing of C. joyneri (BMNH 1924.12.15.87) from Amoy, China.

186.—Ochiai, 1959:208; 1963:88, pi. 18.—Chu, 1963:542, fig.

408—Shen, 1967:211 [Kowloon].

Areliscus joyneri.—Jordan and Snyder, 1900:380 [Tokyo].—

Jordan and Starks, 1906a:241 [Tokyo].—Smith and Pope,

1906:498 [Kochi].— Snyder, 1912:441 [Kagoshima].—Jordan,

Tanaka and Snyder, 1913:336 [listed].—Jordan and Hubbs,

1925:302.—Kuroda, 1931:122.—Kamohara, 1938:62 [Tosa].—

Kuronuma, 1939:83.—Yanai, 1950:22.—Kuroda, 1951:390.—

Kamohara, 1952:85.—Honma, 1952:225.—Chen and Weng,

1965:100, fig. 70 [Tung Kong].

Cynoglossus (Areliscus) lighti Norman, 1925:270 [type-locality:

Amoy and Wenchow].

Cynoglossus lighti—Chu, 1931:95.—Wu, 1932:155 [TangTchou and Amoy].—Chabanaud, 1951a:270.—Wang,

1933:63, fig. 30.

Trulla lighti.—Fowler, 1934b:216.—Chen, 1951:123.

Areliscus tenuis Oshima, 1927:201 [type-locality: Tainan].

Cynoglossus tshusanensis Chabanaud, 1951d:270 [type-locality:

Tshusan Archipelago, China].

DESCRIPTION.—Based on 10 specimens, 193.0-

199.0 mm SL.

Depth of body 20.28-43.64 (M = 25.88), length of

head 18.86-40.00 (M = 23.75) percent of standard

length. Diameter of eye 7.14-10.14 (M = 8.80), interorbital space 4.76-7.95 (M = 4.46) percent of length

of head. Two nostrils on ocular side, anterior nostril tubular in front of lower eye, posterior nostril

simple in anterior half of interorbital space. Snout

obtusely pointed, 32.73-42.03 (M = 36.54) percent

of length of head, rostral hook rather short and

extending to front of anterior nostril. Maxillary

extending well beyond posterior border of fixed eye;

angle of mouth extending to almost below vertical

from posterior half of fixed eye or even beyond,

being more or less midway between tip of snout and

branchial opening; tip of snout to angle of mouth

47.62-60.32 (M = 51.34), angle of mouth to branchial opening 47.06-63.49 (M = 52.26) percent of

length of head.

Scales: Ctenoid on ocular side, including those

of lateral lines; on the blind side scales are cycloid

on head, weakly ctenoid on body.

Lateral-Line System: Usually three lateral lines

on eyed side, the dorsolateral line undulates and

runs backward along dorsal contour of body, usually

entering dorsal fin along 7th ray counted from the

rear, midlateral line with 66-72 (M •» 70) scales,

11-12 (M = 12) scales between middle and upper

lateral line, ventrolateral line usually present, entering anal fin along 5th to 7th ray counted from

the rear. No lateral line on blind side.

Interlinear scale rows II 12

Frequencies 1 8

Fins: Dorsal with 105-112 (M = 109) rays, anal

with 81-85 (M = 83) rays, caudal 10 in 7 specimens

(radiographs).

Vertebrae: 50-53 (M = 51) comprising 9 abdominal and 41—44 caudal elements in 7 specimens

(radiograph).

Coloration: Upper side brownish, blind side

yellowish white in preserved specimens.

Size: Largest specimen examined, 231.5 mm SL,

is from Japan (BMNH 1878.4.5.94).

DISTRIBUTION.—From South China Sea through

Taiwan, Tung-hai, Yellow Sea, Po-hai to Korea and

Japan.

DIAGNOSIS AND AFFINITIES.—Cynoglossus joyneri

is closely related to C. interruptus; it is, however,

distinguished by its longer obtusely pointed snout

(36.54 percent cf. 31.81 percent of head length) and

the situation of the angle of mouth, which is somewhat midway between the tip of snout and branchial

opening, whereas it lies very much nearer to the tip

of snout than to branchial opening in the case of

C. interruptus.
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

NOTE ON SYNONYMY.—Gunther (1878:486) described C. joyneri on the basis of two specimens,

236 and 249 mm in TL, from Tokei, Japan and

characterized it as having three lateral lines, the

upper and the middle being separated by 13 scale

rows. Norman (1925:270) described a similar species C. lighti on the basis of four specimens, 107

to 160 mm in TL, from Amoy and Wenchow and

differentiated it from C. joyneri by its slender body

and shorter head. Comparison of a number of specimens from Japan has clearly indicated that the

differences noted by Norman are not significant to

retain C. lighti as a distinct species from C. joyneri.

Oshima (1927:201) and Chabanaud (1951d:270)

described C. tenuis and C. tshusanensis from Tainan

and Tshusan Archipelago, China, respectively.

Oshima's description was based on a single specimen, 170 mm in TL, and he characterized it as

having three lateral lines, the upper and the

middle being separated by 12 rows of scales, and

considered it "closely related to Areliscus joyneri

differing from it in having a lower body and

ctenoid scales on both sides." Cynoglossus joyneri

too has ctenoid scales on both sides except scales

of the head of the blind side, which are cycloid.

Ochiai (1963:90) synonymized C. tenuis with C.

lighti, which is a junior synonym of C. joyneri.

Cynoglossus tshusanensis was described on the basis

of three specimens, 81.5 to 97 mm in TL, characterized as with two nostrils, contiguous eyes, and

three lateral lines on ocular side, the middle and

the upper being separated by 11-12 scales. I have

examined the types of C. tshusanensis in London

and compared them with the types of C. joyneri;

the differences noted between them were found

only to be attributable to intraspecific variability

and hence they are synonymized here.

TYPE SPECIMENS.—1 specimen, BMNH 1878.4.-

15.94, 231.5 mm SL, lectotype of C. joyneri, designated by Chabanaud (1951d:269), Jokee, Japan,

coll. Joyner. 1, BMNH 1878.4.15.95, 229 mm SL,

paralectotype of C. joyneri, designated by Chabanaud, obtained along with the lectotype. 1, BMNH

1924.12.15.87, 131 mm SL, lectotype of C. lighti,

Amoy, China, coll. Light. 2, BMNH 1924.12.15.88-

89, 131, and 169 mm SL, paralectotypes of C.

lighti, collected along with the lectotype. 1,

BMNH 1924.12.15.90, 117 mm SL, paralectotype

of C. lighti, Wencho, China, coll. Light. 1, BMNH

1892.12.12.32, 88 mm SL, holotype of C. tshusanensis, Chusan Is., China, coll. Walker. 2, BMNH

1892.12.12.33-34, 74 and 77 mm in SL, paratype of

C. tshusanensis, collected along with the holotype.

OTHER MATERIAL EXAMINED.—JAPAN: 3 specimens, USNM

6122, E coast of Japan, coll. Blake. 1, USNM 59763, Kochi,

Japan, coll, H. M. Smith. 1, USNM 151796, Shizuoka, Japan,

coll. D. S. Jordan, 1922. 2, KU 12545 and 12559, Yamaguchi,

Japan. 2, KU 26965 and 26972, Omuta, Fukuoka Pref.,

Japan.

The itinus complex

Cynoglossus itinus, the only species of the itinus

complex, has its center of distribution in Japan

and is related to the kopsi complex, both having

descended from a common ancestral stock. The

principal character distinguishing this species is

the absence of the posterior nostril.

14. Cynoglossus itinus (Snyder)

PLATE 7

Trulla itina Snyder, 1909:609 [type-locality: Naha, Okinawa,

Japan].—Jordan, Tanaka, and Snyder, 1913:337.—Kamohara, 1952:58.—Kuroda. 1954:66.

Cynoglossus itinus.—-Ochiai, 1959:200; 1963:79, pi. 17: fig. 1.

Cynoglossus punctatus Shen, 1969:21. figs. 9-19 [type-locality:

Hong Kong].

DESCRIPTION.—Based on 8 specimens, 99.5-131.0

mmSL.

Depth of body 25.40-28.64 (M = 27.07), length

of head 19.44-21.61 (M = 20.56) percent of standard

length. Diameter of eye 10.87-15.38 (M = 13.22),

interorbital space 2.08-7.14 (M = 4.35) percent of

length of head. Anterior nostril on ocular side

tubular, in front of lower eye, posterior nostril absent. Snout rounded, 25.0-30.43 (M = 27.24) percent

of length of head, rostral hook short, not extending to vertical through the front of anterior nostril.

Maxillary extending to below posterior half of

fixed eye; angle of mouth extending to below vertical from middle of fixed eye, much nearer to tip

of snout than to branchial opening, tip of snout to

angle of mouth 36.54-45.24 (M = 40.55) angle of

mouth to branchial opening 55.77-69.05 (M =

61.34) percent of length of head.

Scales: Ctenoid on ocular side including those

on lateral lines; on blind side, cycloid on head and

weakly ctenoid on body.
NUMBER 238 49

Lateral-Line System.—Three lateral lines on ocular side, dorsolateral line slightly undulated, runs

backward along the dorsal contour of body, entering dorsal fin usually along 20th dorsal ray counted

from the rear, midlateral line with 71-78 (M = 74)

scales, 12-14 (M = 13) scales between them; ventrolateral line present. No lateral line on blind side.

Interlinear scale rows 12 13 14

Frequencies 4 3 1

Fins: Dorsal with 102-103 (M = 102) rays, anal

with 83-86 (M = 84) rays, caudal 8 in 3 specimens

(radiographs).

Vertebrae: 50-52, comprising abdominal and

41-43 caudal elements in 3 specimens (radiographs).

Coloration: Upper side light brown with blackish

blotches, blind side yellowish in preserved specimens.

Size: Largest specimen examined, 141 (131 + 10)

mm, is from Miya Aichi, Japan, KU 16057.

DISTRIBUTION.—Japan and Hong Kong.

DIAGNOSIS AND AFFINITIES.—Cynoglossus itinus

has close affinity with C. interruptus and C. joyneri.

It is distinguished, however, by the lack of the

posterior nostril and a reduction in the number of

caudal fin rays.

NOTE ON SYNONYMY.—Snyder (1909:609) described C. itinus on the basis of a single specimen,

115 mm in TL, from Naha, Okinawa, Japan and

characterized it as having a single nostril and three

lateral lines on ocular side, the upper and the

middle being separated by 12 or 13 scale rows. Recently, Shen (1969:21) described C. punctatus from

seven specimens, 110 to 136.5 mm SL, from Tolo

Harbor, Hong Kong. In having three lateral lines,

contiguous eyes, only one nostril, and the ctenoid

nature of the scales, C. punctatus exhibits the

same features as the present species.

TYPE SPECIMEN.—Holotype of C. itinus, USNM

62957, 105 mm SL, Okinawa, Japan, coll. Albatross.

OTHER MATERIAL EXAMINED.—JAPAN: 1 specimen, USNM

71610, Satsuma, coll. Bureau of Com. Fisheries. 2, USNM

72022, Satsuma, coll. Bureau of Com. Fisheries. 2,

USNM 72023, Tanegashima, coll. Albatross. 1, KU 15351,

Owase, coll. Dr. T. Iwai, 15.1.1950. 1. KU 16057, Miya

Aichi. 1, MCZ 31166, Japan, coll. E. L. Morse.

The ogilbyi complex

The center of distribution of the ogilbyi complex

is in Australia. Like the itinus complex, it is also

closely related to the kopsi complex by the nature

of the eyes situated close together, with a very narrow interorbital space, and the variable nature of

the lateral-line system. Both C. itinus and C. ogilbyi complexes have evolved (the former in Japanese

waters and the latter in the Australian waters} from

a common ancestral stock that spread northward

and southward from the generic evolutionary center. The most outstanding characteristics of the

members are: contiguous or closely situated eyes

with a narrow interorbital space, two or three (C.

maculipinnis) lateral lines, the dorsolateral line

extending full extent or interrupted, and large

scales, the interlinear scale count being 8 or 11 to

14 and usually eight rays in caudal fin and cycloid

scales on the blind side. Included in this complex

are C. ogilbyi, C. maculipinnis, and C. broadhursti.

15. Cynoglossus ogilbyi Norman

Cynoglossus ogilbyi Norman, 1926:304, fig. 14 [type-locality:

southern Queensland].

DESCRIPTION.—Based on the holotype of C. ogilbyi, 179.7 mm SL.

Depth of body 27.88, length of head 18.92 percent of standard length. Diameter of eye 12.65,

interorbital space narrow, 2.06 percent of length

of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior nostril

simple, just in front of interorbital space. Snout

rounded, 26.47 percent of length of head, rostral

hook short, extending to just in front of anterior

nostril. Maxillary extending beyond posterior border of fixed eye; angle of mouth extending to below vertical from posterior half of fixed eye, much

nearer to tip of snout than to branchial opening;

tip of snout to angle of mouth 40.0, angle of mouth

to branchial opening 58.53 percent of length of

head.

Scales: Ctenoid on ocular side, cycloid on blind

side.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line incomplete, extending along

the dorsal contour of the body and extending the

dorsal fin at 31st dorsal ray, counted from the rear,

midlateral line with 66 scales and 8 scales between

the middle and upper lateral lines. No lateral line

on blind side.

Fins: Dorsal with 115 rays, anal with 95 rays,

caudal 8 in 1 specimen (radiograph).
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vertebrae: 56, comprising 9 abdominal and 47

caudal elements in 1 specimen (radiograph).

Size: Only specimen examined, 186 (179 + 7) mm,

is the holotype.

DISTRIBUTION.—Southern Queensland.

DIAGNOSIS AND AFFINITIES.—The nearest relative

of C. ogilbyi is C. maculipinnis; it differs from C.

maculipinnis in having larger scales (interlinear

scale row count 8 cf. 10-13, M = 11) and cycloid

scales on the blind side.

TYPE SPECIMENS.—Holotype of C. ogilbyi, AMSE 2796, 179.7 mm SL, southern Queensland.

16. Cynoglossus maculipinnis Rendahl

PLATE 7

Cynoglossus maculipinnis Rendahl, 1921:17 [type locality:

northwest Australia].—Norman, 1926:302.

Cynoglossus maccullochi Norman, 1926:302, fig. 13 [typelocality: 7-10 miles northwest of Hummocky Is., east coast

of Queensland].

DESCRIPTION.—Based on 11 specimens, 42.5-

139.0 mm SL.

Depth of body 23.89-29.31 (M = 26.58), length of

head 19.0-24.04 (M = 21.42) percent of standard

length. Diameter of eye 11.90-20.00 (15.79), very

narrow, 1.59-5.26 (M = 2.97) percent of length of

head. Two nostrils on ocular side, anterior nostril

tubular, in front of lower eye, posterior nostril in

anterior part of the interorbital space, or immediately in front of the eyes. Snout rounded, 28.00-

35.00 (M = 31.16) percent of length of head; rostral

hook short, ending in front of anterior nostril.

Maxillary extending to below posterior half of

fixed eye; angle of mouth extending to below vertical from middle of fixed eye, nearer to tip of

snout than to branchial opening; tip of snout to

angle of mouth 41.46-50.00 (M = 44.75), angle of

mouth to branchial opening 50.00-60.00 (M =

56.33) percent of length of head.

Scales: Moderate in size, ctenoid on both sides.

Lateral-Line System: Usually two lateral lines on

ocular side, occasionally three or rarely one; dorsolateral line extends to the full extent, very rarely

ending at two-thirds of body;

 8 midlateral line with

64-76 (M = 70) scales, 10-13 (M = 12) scales

* In one of the specimens examined from Lindeman Island,

the dorsolateral line is absent, but the midlateral line is

present.

between middle and upper lateral lines. No lateral

line on blind side.

Interlinear scale rows 10 11 12 13

Frequencies 1 2 6 1

Fins: Dorsal with 102-114 (M = 107) rays, anal

with 79-89 (M = 82) rays, caudal 8 (10) in 7 specimens (radiographs).

Vertebrae: 48-53, comprising 9 abdominal and

39-44 caudal elements in 7 specimens (radiographs).

Coloration: Upper side brownish, with darker

patches forming six or seven irregular bands; two

black dots on caudal fin; lower whitish or yellowish

in preserved specimens.

Size: Largest specimen examined, 150 (139 +11)

mm, is obtained off Lindeman Island, AMS 1A

6810.

DISTRIBUTION.—Australia: Western, Northern

Territory, and Queensland; recorded up to a depth

of 72 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus maculipinnis is closely allied to C. ogilbyi and C. broadhursti. The three species, forming a species-complex,

seem to have evolved in Australia from a common

ancestral stock. The differences between C. maculipinnis and C. ogilbyi have been outlined under C.

ogilbyi; from C. broadhursti it differs in having

ctenoid scales on both sides of body (C. broadhursti,

like C. ogilbyi, has cycloid scales on blind side).

NOTE ON SYNONYMY.—Rendahl (1921:17) described C. maculipinnis based on a single specimen,

from northwestern Australia, and characterized it

with two lateral lines on the ocular side separated

by 13 rows of scales, ctenoid scales on both sides of

the body, two nostrils, the upper eye situated forward by a third of the eye diameter and the interorbital space less than the diameter of eyes.

Norman (1926:302) described C. maccullochi on

the basis of a single specimen, 190 mm in TL, from

Hummocy Island on the east coast of Queensland

collected by Endeavour, and related it to C. maculipinnis; C. maccullochi, however, had larger eyes

and three lateral lines on ocular side. In the

British Museum there is a specimen, BMNH

1925.7.22.84, 107.5 mm SL, collected by Endeavour

from the Moreton Bay on the east coast of Queensland that has only two lateral lines, the upper and

the middle lines being separated by 13 rows of

scales. In another specimen from Lindeman Island

(QMI 15454), 110.5 mm SL, only one lateral line,
NUMBER 238 51

the midlateral line, is present while four other

specimens, QMI 15454, exhibit two lateral lines.

Comparison of a number of specimens has revealed

that the larger eye and an additional lower lateral

line noted by Norman are attributable to intraspecific variation in the species, and C. maculipinnis,

having priority over C. maccullochi, is used here

as the valid name for the species.

TYPE SPECIMEN.—Holotype of C. maccullochi,

AMS 2693, Hummocky Island, east coast of Queensland, coll. Endeavour, examined by Dr. John R.

Paxton on my behalf.

OTHER MATERIAL EXAMINED.—1 specimen, AMS IB 3306,

Repulse Bay, Queensland, 1954. 1, AMS 1A 6810, off Shaw

and Lindeman Is. 1, WAM P 11729 Exmouth Gulf, W

Australia, coll. R. McKay. 1, WAM P 11733. Exmouth

Gulf. 1, WAM P 11799, Careening Bay, Garden Island, W

Australia, coll. M. Graham, 20.1.63. 5, QM 1.15452-56,

Lindeman Is. 1, MCZ 38506, Moreton Bay, S Queensland,

coll. E. J. Coulter; 6.6.52. 1, USNM 174029. Port Bradshaw,

Australia, N T, coll. R. R. Miller, 25.7.48. 1, BMNH

1925.7.22.84, Moreton Bay, S Queensland, coll. Endeavour. 1,

BMNH 1933.8.14.24, NW Australia, coll. Gray. 2. BMNH

1933.8.12.56-57. NE of Low Island, Great Barrier Reef, coll.

G. B. R. Expd.

17. Cynoglossus broadhursti Waite

FIGURE 23; PLATE 8

Cynoglossus broadhursti Waite, 1905:73. pi. 8; fig. 2 [typelocality: off Carnarvon to the northward of Houtman's

Abrolhos].—Norman, 1926:302 [mouth of Murray R.].

DESCRIPTION.—Based on 4 specimens, 183.0-229.0

mm SL.

Depth of body 26.78-29.26 (M = 27.94), length of

head 17.72-20.74 (M = 19.00) percent of standard

length. Diameter of eye 10.47-11.84 (M = 11.15), interorbital space 7.37-10.29 (M = 8.43) percent of

length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior nostril simple, in the anterior half of the

interorbital space. Snout rounded, 26.47-34.21

(M = 30.36) percent of length of head, rostral hook

short, extending to front of the anterior nostril.

Maxillary extending to below posterior half of

fixed eye; angle of mouth extending to below vertical from middle of fixed eye, nearer to tip of

snout than to branchial opening; tip of snout to

angle of mouth 40.00-47.37 (M = 43.48), angle of

mouth to branchial opening 52.63-55.81 (M =

54.53) percent of length of head.

Scales: Ctenoid on ocular side except those of

lateral lines; scales of the blind side and those of

lateral lines on ocular side cycloid.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line running backward along the

dorsal contour of body and entering dorsal fin

along the 6th to 14th ray, counted from the rear,

midlateral line with 72-77 scales, 12-14 scales between middle and upper lateral lines. No lateral

line on blind side.

Interlinear scale rows 12 13 14

Frequencies 2 1 1

Fins: Dorsal with 105-110 (M = 107) rays, anal

with 83-87 (M = 85) rays, caudal 10 in 5 specimens

(radiographs).

Vertebrae: 51-54 comprising 9 abdominal and

42-45 caudal elements in 5 specimens (radiograph).

Coloration: Upper side uniformly brownish,

lower whitish or yellowish in preserved specimens.

DISTRIBUTION.—Western and southern Australia.

Size: The largest specimen examined, 245 (229

+ 16) mm, is from the mouth of the Murray River.

DIAGNOSIS AND AFFINITIES: Cynoglossus broadhursti is readily distinguished from its close relatives, C. maculipinnis and C. ogilbyi, by a greater

interlinear scale count (12-14, M = 12 cf. 8 in C.

ogilbyi and 10-13, M = 11 in C. maculipinnis).

TYPE SPECIMEN.—Holotype of C. broadhursti, a

specimen trawled off Carnarvon to the northward

of Houtman's Abrolhos, in Australian Museum,

Sydney (not examined).

OTHER MATERIAL EXAMINED.—2 specimens, AMS 1.10360 and

1,10359, mouth of Murray River, S Australia. 2, WAM P

14739 and P 14741. Shark Bay, W. Australia, coll. W. and

W. Poob Bros., 18.9.65. 1, BMNH 1925.722.83. W Australia,

coll. Endeavour.

The ecaudatus complex

Cynoglossus ecaudatus and its relatives C. sinusarabici, C. dollfusi, and C. cadenati have their

center of distribution in the western sector of the

Indian Ocean, including the Red Sea and Gulf of

Suez, the eastern coast of the Mediterranean Sea

and the Senegal-Ghana coast. This complex is related to the kopsi complex and is most probably

descended from a common ancestral stock.

The principal characters distinguishing this complex are the large contiguous eyes with a narrow
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Icm

FIGURE 23.—Outline drawing of C. broadhursti (BMNH 1925.7.22.83) from Western Australia.

interorbital space, one or two lateral lines, and the

ctenoid scales on both sides of the body.

18. Cynoglossus ecaudatus Gilchrist

FIGURE 24; PLATE 8

Cynoglossus ecaudatus Gilchrist, 1908:162, pi. 46 [typelocality: off Tugela River].—Gilchrist and Thompson,

1917:398.—Thompson, 1918:128.—Regan. 1920:222 [Natal]—

Bonde, 1922:25.

Areliscus ecaudatus.—Barnard, 1925:415 [Natal and Zululand

coast].

Cynoglossoides ecaudatus.—Smith, 1949:165. pi. 10.

Cynoglossus kopsii [not Bleeker].—Regan, 1908:235 [Seychelles].

Areliscus natalensis Bonde, 1922:23, pi. 4; fig. 2 [type-locality:

Natal Seas, South Africa].

Cynoglossus hunteri Bonde, 1925:293 [type-locality: Delagoa

Bay].—Barnard, 1925:413 [Delagoa Bay]; 1937:41.

Cynoglossus lingua [not Ham Buch.].—Smith, 1949:166

[Delagoa Bay].

Cynoglossus brachycephalus [Cynoglossus seychellensis

Chabanaud (in Schedule)], 1951b:78 [type-locality: Seychelles].

Cynoglossus kopsi diagramata Chabanaud [in part], 1951d:273

[type-locality: Seychelles].

DESCRIPTION.—Based on 4 specimens, 101.5-150.0

mm SL, including the holotype of C. natalensis

Bonde.

Depth of body 25.36-28.02 (M = 27.00), length

of head 19.70-21.74 (M = 20.32) percent of standard

length. Diameter of eye 12.50-13.33 (M = 13.05)

percent of length of head, interorbital space absent.

Two nostrils on ocular side, anterior nostril tubular in front of lower eye, posterior nostril simple,

immediately in front of the eyes. Snout obtusely

rounded, 22.50-33.33 (M = 26.31) percent of length

of head; rostral hook short, ending in front of anterior nostril. Maxillary extending to below posterior half of fixed eye; angle of mouth extending

to below vertical from middle of fixed eye, mudi

nearer to tip of snout than to branchial opening,

tip of snout to angle of mouth 34.78-45.00 (M

= 39.95), angle of mouth to branchial opening

54.35-65.00 (M = 58.59) percent of length of head.

Scales: Ctenoid on both sides, including those on

lateral lines.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line ending at a point between

middle and two-thirds of the length of the body,

midlateral line with 64-66 scales, 10-12 scales between middle and upper lateral line. No lateral

line on blind side.

Interlinear scale rows 10 12

Frequencies 2 2

Fins: Dorsal with 108-110 rays, anal with 86-87,

caudal 8 in 2 specimens (radiographs).

Vertebrae: 52-53, comprising abdominal and

43-44 caudal elements, in two examples in 2 specimens (radiographs).

Coloration: Upper side brown, with more or less

distinct dark crossbands, lower whitish in preserved

specimens.

Size: Largest specimen examined, 166 mm, is

from Somali coast.

DISTRIBUTION.—All along East Coast of Africa

and Seychelles; recorded from 26-000 fins.

DIAGNOSIS AND AFFINITIES.—Superficially and in

several morphometric and meristic characters C

ecaudatus is more closely allied to C. cadenati of

the Senegal-Ghana species than to C. sinusarabici or

C. dollfusi. From C. cadenati it can, however, be

distinguished by the fewer number of rays in the
NUMBER 238 53

caudal fin (8 cf. 10). It is quite conceivable that

these two species have descended from a common

ancestral stock that spread to west Africa through

the Mediterranean during comparatively recent

times when marine connections between the Red

Sea and the Mediterranean existed.

NOTE ON SYNONYMY.—Gilchrist (1908:162) described C. ecaudatus (spelled acaudatus) on the

basis of a single specimen, about 100 mm SL obtained off Tugela River, and characterized it as

having contiguous eyes, ctenoid scales on both sides,

and two lateral lines on ocular side separated by

10 rows of scales. I have examined the holotype of

C. ecaudatus in Cape Town and found traces of

the lower lateral line along the abdominal region.

Cynoglossus natalensis, described on the basis of a

single specimen, 125 mm in TL from Natal and

characterized as having contiguous eyes, ctenoid

scales, and three lateral lines, the lower one being

faintly represented just behind the operculum only

for an inch, is the same species as C. ecaudatus. I

have examined the holotype of C. natalensis in

London and found it identical with C. ecaudatus.

Bonde (1925:293) described C. hunteri on the

basis of a single specimen, 75 mm in TL, from

Delagoa Bay and related it to C. lingua. The type

is not traceable in the South African Museum, but

from the description it does not appear to differ in

any significant way from C. ecaudatus. I have,

therefore, synonymized it with C. ecaudatus.

TYPE SPECIMENS.—Holotype of C. ecaudatus in

South African Museum, off Tugela River. Holotype

of C. natalensis, BMNH 1922.3.27.18, 116 mm SL,

Natal, 29 fms, coll. S.S. Pickle.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH

1908.3.23.148 (one of the paratypes of kopsi diagramata

Chabanaud), Amirantes, Seychelles, coll. Gardiner. 1, BMNH

1969.5.14.401, Seychelles, coll. Wright. 1, BMNH 1904.11.4.5,

S Africa, coll. J. D. F. Gilchrist, 2, SOSC Ref 145, Somali

coast, 76-80 fms, 17.2.64.

19. Cynoglossus cadenati Chabanaud

PLATE 9

Cynoglossus cadenati Chabanaud, 1947a:441 [type-locality:

off Senegal coast, west Africa]; 1949b:205.—Cadenat,

1960:1383 [Accra and Sierra Leone].

Cynoglossus cadenati cadenati.—Chabanaud, 1949b:206 [typelocality: Senegal].

Cynoglossus cadenati honoris Chabanaud, 1949b:206 [typelocality: coasts of Sierra Leone].

Holotype: A specimen of 145.0 mm SL, off Senegal

coast, west Africa, MNHP 49-20.

DESCRIPTION.—Based on 4 specimens, 104-159

mm SL, including the holotype and the paratype of

C. cadenati and also the holotype of C. cadenati

honoris Chabanaud.

Depth of body 18.99-23.00 (M = 20.71), length

of head 18.60-21.15 (M = 19.67) percent of standard

length. Diameter of eye 13.33-16.67 (M = 14.34),

interorbital space 2.38-9.09 (M = 5.99) percent of

length of head. Two nostrils on ocular side, anterior

nostril tubular in front of lower eye, posterior

nostril simple, in the anterior half of the interorbital

space. Snout rounded, 28.81-31.00 (M = 28.51)

percent of length of head, rostral hook short, extending to front of anterior nostril. Maxillary

extending to below posterior border of fixed eye;

angle of mouth extending to below vertical from

posterior half of fixed eye; angle of mouth reaching

below posterior half of fixed eye, nearer to tip of

snout than to branchial opening; tip of snout to

FIGURE 24.—Outline drawing of C. ecaudatus (BMNH 1922.3.27.18) from Natal.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

angle of mouth 40.48-45.83 (M = 43.31), angle of

mouth to branchial opening 52.08-57.14 (M =

54.68) percent of length of head.

Scales: Ctenoid on both sides, including those of

lateral lines.

Lateral-Line System: Two lateral lines on ocular

side, midlateral line with 68-72 (M = 70) scales

and 11-12 scales between them. No lateral line on

blind side.

Interlinear scale rows 11 12

Frequendes 2 2

Fins: Dorsal with 109-115 (M = 112) rays, anal

with 87-88 (M = 87) rays, caudal (9) 10 in 3 specimens (radiographs).

Vertebrae: 52-53, comprising abdominal and 43-

44 caudal elements in 3 specimens (radiographs).

Coloration: Upper side uniformly brown, lower

whitish in preserved specimens.

Size: Largest specimen examined, 173 mm, is from

Liberia, off St. Paul River.

DISTRIBUTION.—West Africa (Senegal to Ghana),

recorded from a depth of 6-12 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus cadenati

is closely related to both C. ecaudatus and C.

dollfusi. In the number of midlateral scale rows,

anal and dorsal fin ray counts, and the vertebral

number these three species are very closely allied.

Cynoglossus cadenati, however, differs from both

species in higher caudal fin ray count of 10, whereas

in the other species there are only 8.

The close affinity of these three species is of

considerable zoogeographical significance. It is conceivable that they evolved from a common ancestral

stock in comparatively recent times when a probable

connection between the Red Sea and the Mediterranean existed, which zoogeographically must not

be earlier than early Pleistocene.

NOTE ON SYNONYMY.—Chabanaud (1947a:441)

described C. cadenati on the basis of three specimens, 106-159 mm in TL, from Senegal and Sierra

Leone, west Africa and characterized it as having

two lateral lines on ocular side separated by 11-12

scale rows and related it to C. canariensis. Cynoglossus cadenati honoris was rightly included in the

synonymy of C. cadenati by Chabanaud (1955a:450).

TYPE SPECIMENS.—MN HP 49.20, 145 mm SL,

holotype of C. cadenati, Senegal, W Africa, coll.

M. J. Cadenat, 1947. 1, MNHP B 2547, 104 mm

SL, paratypes of C. cadenati, collected along with

the holotype. MNHP 49.21, 106.5 mm SL, holotype

of C. cadenati honoris, Sierra Leone, J. Cadenat,

1947.

20. Cynoglossus dollfusi (Chabanaud)

Paraplagusia dollfusi Chabanaud, 1931:303 [type-locality:

Suez Canal].

Cynoglossus dollfusi.—Gruvel and Chabanaud, 1937:8 [Suez

Canal].

Trulla dollfusi.—Fowler. 1956:183.

Cynoglossus lingua [not Hamilton-Buchanan].—Gruvel and

Chabanaud, 1937:10 [Suez Canal].

Cynoglossus cleopatrides Chabanaud, 1949e:146 [type-locality:

Suez Canal].

DESCRIPTION.—Based on the holotype of C.

cleopatrides Chabanaud, 129.0 mm SL.

Depth of body 18.99, length of head 18.60 percent

of standard length. Diameter of eye 16.67, interorbital space 8.33 percent of length of head. Two

nostrils on ocular side, anterior nostril in front

of lower eye, posterior nostril in the anterior half of

the interorbital space. Snout rounded, 27.08 percent

of length of head, rostral hook rather short, ending

in front of the perpendicular of the anterior border

of fixed eye. Maxillary extending to below posterior

half of fixed eye; angle of mouth extending to

below vertical from middle of fixed eye, nearer to

tip of snout than to branchial opening, tip of snout

to angle of mouth 45.83, angle of mouth to

branchial opening 52.08 percent of length of head.

Scales: Moderate size, ctenoid on ocular side,

including those on lateral lines, weakly ctenoid on

blind side.

Lateral-Line System: Two lateral lines on ocular

side, midlateral line with 70 scales, 11 scales between

middle and upper lateral lines. No lateral line on

blind side.

Fins: Dorsal with 115 rays, anal with 85 rays,

caudal 8 (radiograph).

Vertebrae: 53, comprising 9 abdominal and 44

caudal elements (radiograph).

Coloration: Upper side brownish, lower whitish

or yellowish in preserved specimens.

Size: The only specimen examined is the holotype

of C. cleopatrides.

DISTRIBUTION.—Suez Canal.

DIAGNOSIS AND AFFINITIES.—The nearest relative

of C. dollfusi is C. cadenati; it differs from C.

cadenati in having a lesser number of rays in the

caudal fin.
NUMBER 238 55

NOTE ON SYNONYMY.—Chabanaud described C.

dollfusi from the Suez Canal, basing his description

on a single specimen, 33.0 mm in TL, and characterized it as having three lateral lines in the ocular

side, 11 series of scales between the middle and

upper lateral lines, and with ctenoid scales on both

the sides. The type of C. dollfusi is not available in

Paris but Gruvel and Chabanaud (1937:35, figs.

9-12) redescribed the species based on the same

specimen and illustrated it. I examined the holotype

of C. cleopatrides Chabanaud and found it to conform well in all respects, including the number of

scales between the upper and middle lateral lines,

with the description of C. dollfusi except for the

absence of any trace of the ventral lateral line on

the ocular side. As had been pointed out, there is

considerable variation in the development of the

lower lateral line among individuals of the same

species of Cynoglossus (even of the same size), and

the presence or absence of this line should not be

depended upon for species differentiation. Chabanaud's description of C. cleopatrides is, however,

inaccurate in so far as the nature of scales on the

blind side of the body is concerned—the scales on

the blind side of the holotype are ctenoid and not

cycloid as mentioned in the description. I have,

therefore, no hesitation in synonymizing C. cleopatrides with C. dollfusi.

TYPE SPECIMENS.—MN HP 49.24, 129 mm SL,

holotype of C. cleopatrides, Suez Canal, coll. Prof.

R. P. Dollfus, 1928.

21. Cynoglossus sinusarabici (Chabanaud)

FIGURE 25; PLATE 9

Dollfusichthys sinusarabici Chabanaud, 1931:304 [typelocality: Suez Canal]; 1934a: 158.—Gruvel and Chabanaud,

1937:6, 6gs. 5-8.—Ben-Tuvia. 1953:13; 1963:115; 1966:267.

Cynoglossus sinusarabici.—Chabanaud, 1939:30; 1954:465;

1955b: 166.

Cynoglossus branchycephalus [not Bleeker].—Chabanaud,

1951b: 78 [with one lateral line].

DESCRIPTION.—Based on 2 specimens, 83.0 and

99.0 mm SL, the lectotype and paralectotype of

C. sinusarabici.

Depth of body 23.23-25.30 (M = 24.26), length of

head 19.70-20.48 (M = 20.09) percent of standard

length. Diameter of eye 15.38-17.65 (M = 16.52)

percent of head, interorbital space absent. Two

nostrils on ocular side, anterior nostril tubular, in

front of lower eye, posterior nostril simple, in front

of eyes. Snout rounded, 26.47-30.77 (M = 28.62)

percent of length of head, rostral hook, short, ends

in front of anterior nostril. Maxillary extending

below posterior half of fixed eye; angle of mouth

extending to below vertical from middle of fixed

eye, much nearer to tip of snout than to branchial

opening, tip of snout to angle of mouth 43.59-

47.06 (M = 45.32), angle of mouth to branchial

opening 58.82-61.54 (M = 60.18), percent of length

of head.

Scales: Ctenoid on both sides.

Lateral-Line System: Only midlateral line present

on ocular side with 54-60 scales, 11 scales from base

of dorsal to midlateral line at middle of body. No

lateral line on blind side.

Fins: Dorsal with 99-101 rays, anal with 78-79

rays, caudal 8 in 2 specimens (radiograph).

Vertebrae: 48-50 comprising 9 abdominal and

39-41 caudal elements in 2 specimens (radiograph).

Coloration: Upper side uniform brown, lower

whitish in preserved specimens.

FIGURE 25.—Outline drawing of C. sinusarabici (BMNH 1938.107.11)

from Great Bitter Lake, Suez Canal.
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Size: Largest specimen examined, 107 mm (99

mm SL) is the lectotype of C. sinusarabici.

DISTRIBUTION.—Red Sea, Suez Canal, and eastern

Mediterranean (Israel); recorded from a depth of

not more than 75 fms.

DIAGNOSIS AND AFFINITIES.—In C. sinusarabici

only the midlateral line is present, which readily

distinguishes it from the other members of the

ecaudatus complex.

TYPE SPECIMENS.—Chabanaud (1931:304) described C. sinusarabici on the basis of 24 specimens,

thVmaximum length being 134 mm, from the Gulf

of Suez. 1 specimen, MNHP 1967-600a, 99.0 mm

SL, which is selected here as the lectotype, agrees

well with the original description of the species.

Fourteen other paralectotypes, MNHP 1967-6006,

are available in the Paris Museum. The whereabouts

of the rest of the 9 original specimens are not known

tome.

The sealarki complex

Cynoglossus sealarki, C. microphthalmus, C.

Zanzibarensis, and C. capensis form the sealarki

complex, the characteristics of which are the absence

of the posterior nostril, comparatively smaller eyes

with a very narrow interorbital space, three lateral

lines on ocular side, and relatively small scales on

the body, the interlinear scale count being 10-11 or

14-15 or 17. Their distribution is now the central

and western parts of the Indian Ocean. They seem

to have descended from the same ancestral stock

from which the kopsi complex of species evolved.

22. Cynoglossus sealarki Regan

FIGURE 26; PLATE 10

Cynoglossus sealarki Regan, 1908:235, pi. 26: fig. 1 [typelocality: Saya-de-Malha Bank].—Norman, 1928:195.

DESCRIPTION.—Based on 4 specimens, 164-182 mm

SL, including the lectotype and the paralectotypes

of C. sealarki.

Depth of body 23.08-24.39 (M = 23.50), length

of head 18.45-21.67 (M = 19.99) percent of standard

length. Diameter of eye 9.38-12.82 (M = 11.08) percent of length of head, interorbital space 2.56-3.23

(M = 2.90) percent. Anterior nostril of eyed side

tubular, in front of lower eye, posterior nostril

absent. Snout rounded, 30.77-35.48 (M = 32.87)

percent of head, rostral hook very short, extending

to front of anterior nostril. Maxillary extending to

below middle of fixed eye; angle of mouth extending

to below vertical from anterior half of fixed eye,

and nearer to tip of snout than to branchial opening; tip of snout to angle of mouth 30.77-45.16

(M = 37.92), angle of mouth to branchial opening

42.31-61.29 percent of length of head.

Scales: Ctenoid on ocular side including those

of lateral lines; scales on blind side cycloid

anteriorly and ctenoid posteriorly.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 64-66 scales, 10-11 scales

between middle and upper lateral line in 3 specimens. No lateral line on blind side.

Interlinear scale rows 10 11

Frequencies 1 2

I cm

FIGURE 26.—Outline drawing of C. sealarki, paralectotype (BMNH 1908.3.23.154)

from Saya de-malha.
NUMBER 238 57

Fins: Dorsal with 112-116 (M = 114) rays, anal

with 92-96 rays, caudal 10 in 2 specimens (radiograph).

Vertebrae: 57 comprising 9 abdominal and 48

caudal elements in 2 specimens (radiographs).

Coloration: Upper side uniform brown, lower

whitish in preserved specimens.

Size: Largest specimen examined the lectotype of

C. sealarki, is 190 mm.

DISTRIBUTION.—Saya-de-Malha Bank; recorded

from a depth of over 123 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus sealarki

is closely allied to C. microphthalmus. It can, however, be distinguished by its larger scales, the

interlinear scale count being 10-11, whereas C.

microphthalmus has 14 scale rows between its

middle and upper lateral lines.

NOTE ON SYNONYMY.—Regan (1908:235) described C. sealarki based on four specimens, 172 to

190 mm in TL, from Sayade Malha, over 123 fms.

There has been no question of the identity of the

species.

TYPE SPECIMENS.—From the syntype series a

specimen, BMNH 1908.3.23.153, 182 mm SL, from

Saya-de-Malha Bank, coll. Gardiner, is selected here

as the lectotype of C. sealarki, and 3 specimens,

BMNH 1908.3.23.154-156, 164-180 mm SL, collected along with the lectotype are paralectotypes.

23. Cynoglossus microphthalmus (Bonde)

FIGURE 27

Areliscus microphthalmus Bonde, 1922:24, pi. 4: fig. 3 [typelocality: Natal Seas]; 1925:292 [Delagoa Bay].

Trulla microphthalmus.—Barnard, 1925:416.

DESCRIPTION.—Based on the holotype of C. microphthalmus, 164.0 mm SL.

Depth of body 29.27, length of head 21.34 percent

of standard length. Diameter of eye 11.43, interorbital space 5.71 percent of length of head. Anterior

nostril on ocular side tubular, in front of lower

eye, posterior nostril absent. Snout rounded, 30.00

percent of length of head, rostral hook rather

short, extending to front of anterior nostril. Maxillary extending to below posterior border of fixed

eye; angle of mouth extending to below vertical

from posterior half of fixed eye, nearer to tip of

snout than to branchial opening; tip of snout to

angle of mouth 44.29, angle of mouth to branchial

opening 58.57 percent of length of head.

Scales: Ctenoid on ocular side, including those of

lateral lines; scales on blind side cycloid anteriorly

and weakly ctenoid posteriorly.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 79 scales, 14 scales between

middle and upper lateral lines, ventrolateral line

slightly undulated. No lateral line on blind side.

Fins: Dorsal with 108 rays, anal with 86 rays,

caudal 10.

Vertebrae: (Not counted).

Coloration: Upper side light brown, lower whitish

in preserved specimens.

Size: The only specimen examined, 174.5 mm,

is the holotype of C. microphthalmus.

DISTRIBUTION.—Natal, South Africa; recorded

from a depth of 29 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus microphthalmus is closely allied to C. sealarki, but it can

be easily distinguished by its wider interorbital

space (5.71 percent of length of head cf. 2.90

percent).

FIGURE 27.—Outline drawing of C. microphthalmus, holotype (BMNH 1922.3.27.17) from Natal.
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TYPE SPECIMENS.—Holotype of C. microphthalmus, BMNH 1922.3.27.27, 164 mm SL, Natal, coll.

Gilchrist.

24. Cynoglossus zanzibarensis Norman

FIGURE 28; PLATE 10

Cynoglossus (Trulla) zanzibarensis Norman, 1939:105, fig. 36

[type-locality: Zanzibar, 183-293 m].—Nielsen, 1961:226

[off Durban, 430-595 m]; 1964:132 [off Durban, 230 m].

DESCRIPTION.—Based on 15 specimens, 49.0-172.0

mm SL, including the lectotype and paralectotype

of C. zanzibarensis.

Depth of body 21.78-28.19 (M = 25.60), length of

head 20.72-24.10 (M = 21.98) percent of standard

length. Diameter of eye 8.82-13.33 (M = 11.28),

interorbital space absent or narrow, 0.31-4.76 (M —

2.75) percent of length of head. Anterior nostril on

ocular side tubular, in front of lower eye, posterior

nosrril absent. Snout rounded, 27.27-35.0 (M =

32.55) percent of length of head; rostral hook very

short, extending to front of anterior nostril. Maxillary extending to below posterior half of fixed eye;

angle of mouth extending to below vertical from

middle of fixed eye, nearer to tip of snout than to

branchial opening; tip of snout to angle of mouth

37.33-47.06 (M = 41.93), angle of mouth to

branchial opening 53.85-64.71 (M = 58.15) percent

of length of head.

Scales: Ctenoid on both sides, those of head on

blind side rather weakly ctenoid.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 72-76 scales, 13-15 scales

between them. No lateral line on blind side.

Interlinear scale rows 13 14 15

Frequencies 2 4 4

Fins: Dorsal with 116-121 (M = 119) rays, anal

with 94-99 (M = 96) rays, caudal 10 in 6 specimens

(radiographs).

Vertebrae: 51-58, comprising 9 abdominal and

42-49 caudal elements in 6 specimens (radiographs).

I cm

FICURE 28.— C. zanzibarensis, paralectotype, (BMNH 1939524.1810) from Zanzibar area: a, outline drawing of lateral view; b, details of pelvic fin; pelvic fin of eyed side persists as rudimentary structure.
NUMBER 238 59

Coloration: Upper side brownish, lower whitish

in preserved specimens.

Size: Largest specimen examined, 185 mm, is

from the Arabian Sea, lat. 21°23'N, long. 69°46'£.

DISTRIBUTION.—Durban through Zanzibar to

Kenya.

DIAGNOSIS AND AFFINITIES.—Cynoglossus zanzibarensis is closely related to C. capensis, especially

in the position of the eyes, which are close together

leaving a very narrow or no interorbital space. From

C. capensis it is readily differentiated by the ctenoid

scales on both sides of its body.

NOTE ON SYNONYMY.—Originally described on the

basis of five specimens, 130-177 mm in TL, collected

off Zanzibar by John Murray Expedition. Cynoglossus zanzibarensis is well diagnosed and there

has been no confusion with regard to its identity.

TYPE SPECIMENS.—A specimen, BMNH 1939.5.24.-

1813, 149 mm SL, from Zanzibar, which is an

excellent example of the species, is selected here

as the lectotype of C. zanzibarensis and four specimens, BMNH 1939.5.24.1810-12 and 14, 120-169

mm SL are paralectotypes.

OTHER MATERIAL EXAMINED.—S specimens, SOSC Ref 170,

Kenya coast, lat. 02°56'S, long. 40'28'E. coll. Anton Bruun.

1, SOSC Ref 23, Kenya coast, lat 20°50'S, long. 40°31'E. coll.

Anton Brunn. 2, SOSC Ref 170, Mozambique coast, lat.

25°12'S, long. 34°04'E, coll. Anton Bruun. 5. SOSC Ref 170,

Mozambique coast, coll. Anton Bruun.

25. Cynoglossus capensis (Kaup)

PLATE 11

Trulla capensis Kaup, 1858:109 [type-locality: Cape of Good

Hope].—Thompson, 1918:128.—Bonde, 1925:292 [Delagoa

Bay].—Barnard, 1925:416.—Smith, 1949:166, pi. 2: fig. 342.

Plagusia capensis.—Castelnau, 1861:71.

Cynoglossus capensis.—Gunther, 1862:503.—Boulenger, 1902:

4.—Nielsen, 1964:133 [Cape Barracouda, Agulhas Bank]

Areliscus capensis.—Bonde, 1922:24.

DESCRIPTION.—Based on 6 specimens, 87.0-293.0

mm SL.

Depth of body 23.56-43.70 (M = 29.21), length

of head 18.09-34.03 (M = 21.68) percent of standard

length. Diameter of eye 10.71-12.90 (M = 11.54),

interorbital space absent or very narrow, 1.79-1.89

(M = 11.84) percent of length of head. Anterior

nostril on ocular side, in front of lower eye, posterior

nostril absent. Snout rounded, 28.40-35.85 (M =

32.14) percent of length of head, rostral hook very

short, ends well in front of anterior nostril. Maxillary extending to below posterior half of fixed eye;

angle of mouth reaching below vertical from middle

of fixed eye, nearer to tip of snout than to branchial

opening; tip of snout to angle of mouth 37.14

46.77 (M = 43.20), angle of mouth to branchial

opening 56.79-62.50 (M = 59.65) percent of length

of head.

Scales: Ctenoid on ocular side; scales of blind

side cycloid anteriorly, weakly ctenoid posteriorly.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 98-104 scales, 15-17 scales

between middle and upper lateral lines. No lateral

line on blind side.

Interlinear scale rows 15 16 17

Frequencies 1- 2

Fins: Dorsal with 108-110 (M = 109) rays, anal

with 85-86 (M = 85) rays, caudal 10 in 3 specimens

(radiographs).

Vertebrae: 50-52 comprising 9 abdominal and

41-43 caudal elements, in 3 specimens (radiographs).

Coloration: Upper uniformly brown, lower whitish in preserved specimens.

Size: Largest specimen examined, 314 (293 + 21)

mm, is from False Bay, South Africa.

DISTRIBUTION.—South Africa (from Saldanha Bay

to Natal) recorded from a depth of 10-60 fms.

DIAGNOSIS AND AFFINITIES.—Superficially and in

many morphometric characters C. capensis resembles

C. zanzibarensis. It is distinguished, however, by its

deeper body, the cycloid nature of the scales of

the anterior part of the blind side, and 15 to 17

scale rows between its middle and upper lateral

lines.

TYPE SPECIMENS.—Kaup's (1858:108) brief description of C. capensis from the Cape of Good

Hope was based on a single specimen without indication of its size. Kaup apparently did not retain the

specimen on which his description was based and

I have found no evidence that it is extant.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH

1897.10.18.1, False Bay, coll. Gilchrist. 1, BMNH 1891.932.

Algoa Bay, coll. Leslie. 1, BMNH 1904.11.4.4, Cape Point,

coll. Gilchrist. 1, BMNH 1922.35722, Natal. Gilchrist. 2.

MCZ 11184, Cape of Good Hope, coll. Sayard.

The arel group

This group is more highly specialized for a
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

burrowing habit than the previous groups and is

typified by relatively larger adult size, more elongate

body, total absence of any lateral-line system on

blind side, 10 rays in caudal fin, two lateral lines

on ocular side, relatively small eyes separated by a

narrow interorbital space, two nostrils on ocular

side, somewhat obtusely pointed long snout with

angle of mouth situated nearer to branchial opening

than to tip of snout, and large scales, ctenoid on

ocular side and cycloid on blind side. The obtusely

pointed snout with the consequent placing of the

angle of mouth nearer to branchial openings, more

elongate body, and smaller eyes are further specializations toward a burrowing mode of life.

Included in this group is the arel complex consisting of C. arel, C. robustus, and C. lingua; C. arel

with an interlinear scale formula of seven to nine

scales being closest to the evolutionary stem of

this group. Their combined range covers the whole

of the Malay Archipelago extending north to Japan

and west to India, Pakistan, and the Persian Gulf.

This group, which has certain similarities with the

canariensis and bilineatus complexes, appears to

have evolved in the main generic evolutionary

center as an early offshoot from the stock that gave

rise to the canariensis group.

The arel complex

Cynoglossns arel, C. robustus, and C. lingua form

the arel complex, which is the only complex within

the arel group. The characteristics of the complex

are, therefore, the same as given above for the

group.

26. Cynoglossns arel (Schneider)

FIGURE 29; PLATE 11

Pleuronectes arel Schneider, 1801:159 [type-locality: Tranquebar, east coast of Madras].

Cynoglossus arel.—Norman, 1928:201.—De Silva, 1956:199

[Palk Bay, Ceylon].—Saramma, 1963:76 [Kerala coast].

Cynoglossus melampetala Richardson, 1846:281 [China].—

Whitehead, 1969:218, pi. 29a.

Cynoglossus melampe talus.—Gunther, 1862:496.—Bleeker,

1873:131 [Canton].—Gunther, 1880:55 [Hong Kong].—Sauvage, 1881:107 [Swatow].—Wu, 1932:148.—Fowler, 1934b:221.

—Chu, 1963:540, fig. 406.—Shen, 1967:216.

Plagusia grandisquamis Cantor, 1850:1214 [type-locality: seas

of Penang].

Trulla grandisquamis.—Kaup, 1858:109.

Cynoglossus grandisquamis.—Gunther, 1862:503.—Duncker,

1904:169.—Weber and de Beaufort, 1929:208.

Plagusia potous [not potous Cuvier].—Cantor, 1850:1217

[type-locality: seas of Malay Peninsula and islands].

Plagusia macrolepidota Bleeker, 1851a:415 [type-locality:

Batavia]; 1852:25.

Arelia macrolepidota.—Bleeker, 1859:184.

Cynoglossus macrolepidotus.—Gunther, 1862:496.—Bleeker,

1875:34, pi. 242: fig. 2—Day, 1877:434, pi. 96: fig. 3;

1889:455—Alcock, 1889:288.—Rutter. 1897:89.—Seale,

1910:288.—Jenkins, 1910a:30.—Norman, 1928:202, fig. 18.—

Weber and de Beaufort, 1929:205.—Herre, 1933:5 [Sandakan].—Fowler, 1934b:219; 1937:87.—Okada and Matsubara,

1938:437 [Formosa, Java. Banka].—Herre, 1941:392 [Andaman Is.].—Suvatti, 1950:327.—Herre, 1953:190.—Munro,

1955:265, pi. 51: fig. 770.—Fowler, 1956:186, fig. 101.—De

Silva, 1956:197—Scott, 1959:43.—Kuronuma, 1961:32.—

Saramma, 1963:76—Punpoka, 1964:64.—Pradhan, 1964:458.

—Chen and Weng, 1965:95. fig. 66.—Shen, 1967:215.

Plagusia cantoris Bleeker, 1853a: 153 ["Hindustan and Archipelagus, Indicum" based on 2 stuffed specimens of Cantor's

P. potus from Singapore].—Gunther, 1862:502.—Bleeker,

1875:33.

Plagusia oligolepis Bleeker, 1854b:445.

Arelio olegolepis.—Blecktt, 1859:185.

Cynoglossus oligolepis.—Gunther, 1862:496.—Bleeker, 1875:34,

pi. 242: fig. 3.—Steindachner, 1867:587 [Ning-Pou].—

Bleeker, 1873:134 [Ning Pou].—Day. 1877:433. pi. 95: fig. 4;

1889:455.—Alcock. 1887:280.—Johnstone. 1904:209.—Weber,

1913b:441.

Cantoria pinangenensis Kaup, 1858:106 [based on Plagusia

cantoris Bleeker].

Arelia kaupii Bleeker, 1860a:73.

Cynoglossus kaupii.—Gunther, 1862:497.—Bleeker, 1875:32,

pi. 244: fig. 4.—Weber and de Beaufort, 1929:196.

Cynoglossus elongatus Gunther, 1862:501 [type-locality: East

Indian seas].—Bleeker, 1875:34.

DESCRIPTION.—Based on 26 specimens, 150.0-330.0

mm SL, including the holotype of C. macrolepidotus

and C. oligolepis.

Depth of body 19.94-25.81 (M = 22.73), length

of head 19.69-28.39 (M = 23.99) percent of standard

length. Diameter of eye 6.61-10.85 (M = 8.48), interorbital width 1.75-5.21 (M = 3.43) percent of length

of head. Two nostrils on eyed side, the anterior one

tubular in front of lower eye, posterior simple in

the anterior half of the interorbital space. Snout

obtusely pointed, 20.66-41.89 (M = 35.87) percent

of length of head, rostral hook short, extending to

front of the anterior nostril. Maxillary extending to

beyond the fixed eye; angle of mouth extending

to below vertical from hind border of fixed eye or

little beyond, more or less midway between tip of

snout and branchial opening; tip of snout to angle

of mouth 46.25-55.10 (M = 50.71), angle of mouth
NUMBER 238 61

I cm

FIGURE 29.—Outline drawing of C. arel (BMNH 1848.3.16.193) from Chinese seas.

to branchial opening 46.49-53.49 (M = 49.33) percent of length of head.

Scales: Ctenoid on ocular side, including scales on

lateral line, those on head rather weakly serrated;

scales on blind side cycloid.

Lateral-Line System: Two lateral lines on eyed

side, median lateral line with 56-70 scales, 7-9

scales between them. No lateral line on blind side.

Interlinear scale rows 7 8 9

Frequencies 20 43 10

Fins: Dorsal with 116-130 (M=125) rays, anal

with 85-98 (M = 95) rays, caudal 10 in 42 specimens

(radiographs).

Vertebrae: 50-57 comprising 9 abdominal and

41-48 caudal elements in 42 specimens (radiographs).

Coloration: Upper side more or less uniformly

brownish, lower whitish in preserved material.

Size: Largest specimen examined, 365.0 mm SL,

from Madras, India.

DISTRIBUTION.—From Malay Archipelago, through

seas of India to Persian Gulf and through South

China Sea to Philippines and Taiwan; recorded

from a depth of 5-45 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus arel is

closely related to both C. lingua and C. robustus.

It is distinguished by the lower number of interlinear and midlateral scales, anal and dorsal rays,

and vertebrae than the other two species. Cynoglossus arel appears to be less specialized than C.

robustus for a burrowing mode of life in that it

has a shorter snout (mean length 35.87 percent

of length of head cf. 39.36 percent) and larger

eyes (mean diameter of eye 8.48 percent of length

of head cf. 7.84 percent).

NOTE ON SYNONYMY.—Schneider's (1801:159) description of P. arel is not quite adequate to distinguish it from most other species of Cynoglossus. He

described C. arel on the basis of four dried specimens

from Tranquebar on the east coast of Madras,

India, neither giving any illustration of the species

nor mentioning the size of his specimens. In the

original description, the number of scale rows

between the upper and middle lateral lines was

not indicated, which in fact had created the confusion in the identity of the species. Richardson

(1846:281) described C. melampetala from China,

based on John Reeve's collections of Chinese drawings; Cantor (1850:1214) described Plagusia grandisquamis on the basis of a specimen from Penang;

Bleeker (1851a:415) described P. macrolepidota

from Batavia, and Bleeker (1860a:73) described

Arelia kaupii from Sumatra. Bleeker in his Atlas

(1875:34, pi. 242: fig. 2) redescribed C. macrolepidotus and illustrated it. His description and illustration were so good that subsequent workers identified

this common and widely distributed Indo-West Pacific species with C. macrolepidotus, not suspecting

that the earlier name arel was its senior synonym.

Bleeker (1853a: 153), based on two stuffed specimens

of Cantor's Plagusia potous, 217 and 322 mm SL,

from Singapore, described another species P. cantoris, which he differentiated from all other known

species on the basis of both nostrils being placed

above the upper lip in front of the lower eye. I have

examined the types of P. cantoris; even though varnished over, the position of the nostrils cannot be
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

detected. Likewise, Cantor's P. grandisquamis,

which is differentiated by only one nostril in front

of the lower eye, the absence of the narial openings

between the eyes, and two lateral lines on ocular

side separated by six scale rows, is also represented

in the British Museum by a stuffed specimen. I have

examined it and though the presence of the nostrils

in the varnished skin is not discernible, an interlinear count of seven scales can be determined, a

distinctive characteristic feature of C. arel.

While describing P. oligolepis and A. kaupii,

Bleeker did not compare them with any other

species. From his description it would appear that

he considered P. oligolepis different from C. macrolepidotus on the basis of a narrower interorbital

space, the situation of the angle of the mouth being

somewhat nearer to the gill opening than to the

tip of mouth, and a slightly greater depth of the

body. Cynoglossus kaupii, described on the basis

of a single specimen, was differentiated both in its

original description and in Bleeker's Atlas from

all other species by two lateral lines on both sides.

Gunther (1862) considered C. macrolepidotus, C.

oligolepis, and C. kaupii as distinct species, but

about C. kaupii he remarked that "Pleuronectes

arel Bl. Schn. p. 159, or Arelia Schneider Kaup,

I.e., appears to be closely related to this species." He

described another species, C. elongatus, based on

a specimen of Cantor's Plagusia potous from

Penang, and characterized it as a somewhat elongated fish with the depth of the body being "six

times and two-thirds in the total length." I have

not been able to locate Cantor's stuffed specimen

that formed the basis for Giinther's C. elongatus,

but from the description, "two lateral lines on the

left side, separated by about eight longitudinal

series of scales at their point of greatest distance,"

the identity of the species is unmistakable and is

readily referable to C. arel.

Sixty-one years after the original description of

C. arel, Norman (1928:201) examined the type of

C. arel from the Berlin Museum and resurrected

the species. He further synonymized C. oligolepis

with C. arel but somehow retained C. macrolepidotus as a separate species, differentiating it from C.

arel by its somewhat deeper body. I have examined

a large number of specimens of various sizes including specimens from Madras, the type-locality of

C. arel, and compared them with the specimens

labeled as C. macrolepidotus in the British Museum, including a specimen from Bleeker's collection

(BMNH 1862.6.3.8), and I have come to the definite

conclusion that the differences noted by Norman in

the proportional measurements are attributable to

intraspecific variation and that they are conspecific.

Cynoglossus kaupii, from the original description as

well as from Bleeker's excellent illustration, appears

to exhibit no significant difference from C. arel;

eight or nine scales between the lateral lines on the

left side fairly well confirm the identity of the

species. Plagusia melampetalus Richardson (1846)

is synonymized with C. arel, because from the

characterization of the species in the original description, especially the indication that the two

lateral lines on the left side are separated by seven

series of scales, the identity of the species is unmistakable.

TYPE SPECIMENS.—Holotype of P. arel, ZMB 2481,

207 mm SL, 221 mm TL, Tranquebar, east coast

of Madras. (Dr. C. Karrer kindly examined the

type in the Zoological Museum, Berlin and gave the

necessary details).

Bleeker's description of P. macrolepidota was

based on 7 specimens, 155-240 mm in TL, from

Batavia. In his Atlas (1875:34, pi. 242: fig. 2) he

redescribed and illustrated the species based on

17 specimens, 155-240 mm in TL, and illustrated a

specimen of 240 mm in TL. Among the Bleeker

collection in Leiden there are 6 specimens, cataloged

as RMNH 6785 (pers. comm., Dr. M. Boeseman).

The specimen, 218 mm SL (239 mm TL), that

formed the basis for Bleeker's illustration and has

already been indicated as such by a label in the

bottle by Chabanaud in 1946 is selected here as

the lectotype. RMNH 6785 is now restricted to

the lectotype and the rest of the 5 specimens, 134-

190 mm SL (146-206 mm TL), are paralectotypes

and have been recataloged as RMNH 26209 (Dr.

M. Boeseman examined the types on my behalf and

selected the paralectotypes).

Bleeker's description of P. oligolepis was based on

one specimen, 365 mm in TL, from Batavia. Among

the specimens in the Bleeker collection in Leiden,

there are two specimens cataloged as RMNH

6786 (pers. comm., Dr. M. Boeseman). A specimen

measuring 338 mm SL (365 mm TL) is considered

to be the holotype of P. oligolepis and is retained

as RMNH 6786, the other specimen, 272 mm SL

(293 mm TL), having been removed.

Bleeker's description of A. kaupii was based on
NUMBER 238 63

one specimen, 255 mm in TL, from Benkulen,

Sumatra. Among the Bleeker collection in Leiden

there is a specimen, 232 mm SL (248 mm TL), coll.

J. A. W. Van Ophuysen, 1858, cataloged as RMNH

6783 (pers. comm., Dr. M. Boeseman) and it is

the holotype of A. kaupii.

OTHER MATERIAL EXAMINED.—CHINA: 1 specimen, BMNH

1848.3.16.198, China Sea, coll. Vice Admiral E. Belcher. 2,

BMNH 1879.5.14.79-80, Hong Kong, coll. Challenger. 1,

BMNH 1924.12.15.82, Santuas, China, coll. Light. 4, BMNH

1924.12.15.83-86, Amoy, coll. Light. 1, BMNH 1955.5.18.6, S

China, coll. Lin. 1, USNM 6511, China, coll. Stempson. 3,

USNM 36895, Hong Kong, coll. W. Stempson. 1, USNM

56430, China, coll. Jony. 1, USNM 137951, Hong Kong Mkt.,

coll. Albatross. 3, USNM (unregistered), Hong Kong, Fisheries Research Sta, Hong Kong. 1, MCZ 33194, Hong Kong.

1, ANSP 76713, Tai Po, China, coll. G. C. Herklots. 1, ANSP

77399, Hong Kong, China, coll. G. C. Herklots.

INDONESIA: 1, BMNH 1862.53.2, Batavia, coll. P. Bleeker.

1, BMNH 1931.4.23.48, Batavia Mkt., coll. Hardenberg. 1,

BMNH 191.4.23.49, Cheriban, Java, coll. Hardenberg. 1,

BMNH 191.4.23.50, Kelampis. Madora, coll. Hardenberg.

1, USNM 72749, Java, coll. B. Palmer. 4, UMMZ 154921

(OR.274495), Batavia, coll. Hubbs and Hardenberg. 1, MCZ

OR.274867, Java, Surabaya, coll. Hubbs and Palmer. 1,

MCZ OR.274867, Batavia. 1, NHV OR.276012, Padang, coll.

Dr. K. Paul. 1, UZMK OR.275455, lat. 5°56'S, long. 106°34'E,

coll. Th. Mortensen. 2, UZMK (unregistered) Lampong Bay,

coll. Th. Mortensen. 1, UZMK (unregistered), lat. 6°22'S,

long. 105°44'E, coll. Th. Mortensen, 26.7.22. 1, UZMK (unregistered), Bantam Bay (Sta 105), coll. Th. Mortensen, 5.8.22.

THAILAND: 2, UZMK (unregistered), Gulf of Thailand,

lat. 6°57'N, long. 99°23'E, and lat. 7°00'N, long. 99°21'E,

30-26 m, coll. BM-Thai-Danish Expd., 10.2.66. 5, UZMK

(unregistered), Gulf of Thailand, Jan-Feb 1966. 1, UZMK

(unregistered), Gulf of Thailand. 4, ANSP 76816, Thailand,

coll. R. M. Schauensee. 1, ANSP 87212, Bangkok, coll. R. M.

Schauensee. 3, CAS (GVF), lat. 11°59'24"N, long.

102°U'06"E, coll. Naga Expd. 1, CAS (GVF), lat. 13°90'N,

long. 100°45'E, coll. Naga Expd. 1, CAS (GVF), lat.

10°26'13.5"N, long. 99°15'27"E, coll. Naga Expd., 18-5-60. 2,

GAS (GVF), lat. ^"Sg^S'N, long. 100°15'45wE, coll. Naga

Expd., 13-12-60. 1, GAS (GVF), Gulf of Thailand, lat.

11 °49'N, long. 94"49'E, coll. Thai fishermen, 2.1.1960. 4, CAS

(GVF), Gulf of Thailand, coll. Naga Expd., 22.4.1960. 11,

CAS (GVF), lat. 13°05'50"N, long. 100°26'10'fE, coll. Naga

Expd., 13.12.1960. 1, CAS (GVF), lat. 13°05'50"N, long.

100°26'10"E, coll. Naga Expd., 13.12.1960. 2, CAS (GVF),

lat. HMO'N, long. 100°39'E and lat. 11°51'N, long 100°39'E,

coll. Thai fishermen, 10.1.1961.

SINGAPORE: 8, MCZ 332208, Singapore, coll. Dr. G.

Mead. 1, NHV OR.276012, Singapore, coll. Dr. K. Paul. 1,

BMNH 19337.21.27, Singapore, ex. Raffles Mus.

BURMA: 1, BMNH 1928.3.20.96, Gulf of Martaban, ex.

Indian Museum. 1, ZSI

 F 117 ° , Gulf of Martaban, coll.

F 253 Marine Survey. 1, ZSI —=—, Mergui Mkt.

INDIA: 1, BMNH 1888.11.6.45, E coast of Madras, coll.

Thurshton. 1, BMNH 1867.5.30.12, Madras, coll. F. Day. 1.

BMNH 1869-5-30.12. Madras, coll. F. Day. 2, BMNH

1889.2.1.4058, 59, Madras, coll. F. Day. 1, BMNH

1928.3.20.92-91, Orissa coast ex. Indian Museum. 2.

BMNH 1928.3.20.93-94, R. Hooghly, ex. Indian Museum.

1, BMNH 1928.3.20.95, Andamans, ex. Indian Museum. 1,

BMNH 1928.3.20.97, R. Hooghly, ex. Indian Museum.

1, BMNH 1928.3.20.98, R. Hooghly, ex. Indian Museum. 1,

ZSI 1143 (original of Day's pi. 96: fig. 3), Madras, coll. F.

Day. 1, ZSI 1190, Madras, coll. F. Day. 1, ZSI 1458 (original of Day's pi. 95: fig. 4), Madras, coll, F. Day. 1, ZSI 2716

(original of Day's pi. 90: fig. 5), Madras, coll. F. Day. 1,

ZSI 11588, Sandheads, R. Hooghly, coll. R. M. Daley. 4, ZSI

12111-4, Orissa coast, coll. Marine Survey. 2, ZSI 12215,

Orissa coast, 5 mi S of False Pt., 7 fms, coll. Marine Survey.

1, ZSI 12126, Orissa coast, coll. Marine Survey. 1, ZSI

F 8760

—j— , Orissa coast, coll. Bengal Fisheries Golden Crown.

F 1232

2 ZSI —-—, Keechankuppan, 3 mi S of Nagapatnam, coll.

A. G. K. Menon, 8.2.57. 1, ZSI

 F 2826

, Pun coast, coll. N.

Annandale. 1, ZSI —- , locality unknown, coll. Bengal

F 3572

Fisheries Golden Crown. 3, ZSI —r , locality not known,

F 1648

coll. Bengal Fisheries Golden Crown. 3, ZSI ^ , locality

not known, coll. Bengal Fisheries Golden Crown. 1, ZSI

F 1297

—2— , Veerampatnara, 3 mi S of Pondicherry, coll. A. G. K.

Menon, 22.1256. 1, ZSI

 F g1208

, Sonakuppam ViUage, Cuddalore. coll, A. G. K. Menon, Jan 57. 1, ZSI

 F *210

. Pondicherry Mkt., coll. A. G. K. Menon, 25.12.1956. 1, ZSI

F 1229

Tranquebar coast, Tanjore Dt., Madras, coll. A. G. K. Menon,

29.157. 1, ZSI F 1772. Alibagh, Colaba Dt., Maharastra

F 1772

29.157. 1, ZSI , Alibagh, Colaba Dt., Maharastra

State, coll. K. K. Tiwari, 24.1155. 33, SOSC Ace. 4, lat.

14°52'N, long. 96°39'E, Sta S9a, coll. Anton Bruun, 31.3.63.

127, SOSC Ace. 4, lat. 15°04'N, long. 95°51'E, Sta 41, coll.

Anton Bruun 31.3.63. 1, SOSC Ace. 4, lat. 15°20'N, long.

96°24'E, coll. Anton Bruun, 313.63. 6, SOSC Ace. 4, lat.

15°08'N, long. 94°54'E, Sta 42, coll. Anton Bruun, 1.4.63. 3,

SOSC Ace. 4, lat. 21O52'N, long. 91°36'E, Sta 44, coll. Anton

Bruun, 4.4.63. 5, SOSC Ace. 4, lat. 19o50'N, long. 92°55'E.

coll. Anton Bruun, 5.4.63. 4, SOSC Ace. 4, lat. 19°41'N,

long. 93°08'E, coll. Anton Bruun, 5.4.63. 1, SOSC Ace. 23,

lat. 20°22'N, long. 11°47'E, coll. Anton Bruun, 15.11.63 129,

SOSC Ace. 23, lat. 20°20'N, long. 70°50'E, 98 fms. coll.

Anton Bruun, 15.11.63. 1, SOSC Ace. 23, lat. 21"23'N, long.

69°46'E, Sta 211A, coll. Anton Bruun, 16.11.63. 1, SOSC Ace.

23, lat. 22°03'N, long. 68°19'E, coll. Anton Bruun, 18.11.63.

8, SOSC Ace 23, lat. 22°21'N, long. 68O42'E, Sta 217A, 14

fms, coll. Anton Bruun, 18.11.63. 5, SOSC Ace. 23, lat.

22°45'N, long. 68°24'E, 14 fms, coll. Anton Bruun, 18.11.63.

1, SOSC Ace. 23, lat. 23°16'N, long. 67°48'E, 12 fms. Sta 225A,

coll. Anton Bruun, 19.11.63. 2, SOSC Ace 23, lat. 23°00'N.

long. 68°36'E, coll. Anton Bruun, 19.11.63. 2, SOSC Ace 4,
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lat. 14°52'N, long. 96°39'E, coll. Anton Bruun, 31.11.63. 1,

SOSC Ace. 4, lat. 14852'N, long. 96°24'E, Sta 39A., coll.

Anton Bruun, 31.11.63. 1, SOSC Ace. 23, lat. 21°28'N, long.

59°28'E, coll. Anton Bruun, 4.12.63. 2, SOSC Ace. 23, Royapuram Beach, Madras, lat. 13°O7'N, long. 80°20'E, coll.

An ton Bruun, 13.9.66. 1, SOSC Ace. 23, Ennor fish landing,

Madras, lat. 13°13'N, long. 8O825'E, coll. Anton Bruun,

14.9.66. 5, SOSC. Ref 334, Porto Novo, Madras, lat.

llo30'N, long. 79°50'E, 12 fms, coll. Anton Bruun, 21.9.66.

2, SOSC, Porto Novo, Madras, lat. 11°32'N, long. 79°53'E,

coll. Anton Bruun. 1, SOSC, Pondicherry, lat. 11°55'N,

long. 79°53'E, coll. Anton Bruun. 1, specimen, SOSC,

Veerampatnam, Pondicherry, lat. 11°52'N, long. 39°54'E, coll.

Anton Bruun. 1, SOSC Ace. 23. Palk Strait, Madras, lat.

09°20'N, long. 79° WE, coll. Anton Bruun, 29.9.66. 2, SOSC

Ace. 23, Mandapam Camp, Tamil Nadu, lat. 09°25'N, long.

79°10'E, coll. Anton Bruun, 29.9.66. 3, SOSC Ref. 334, lat.

08°39'N, long. 78°15'E, coll. Anton Bruun, 1.10.66. 1, SOSC,

Porto Novo, Madras, coll. Anton Bruun, 2.10.66. 1, SOSC

Ref. 381, Porto Novo, Madras, coll. Anton Bruun, 9.10.66. 1,

SOSC, Madras, coll. T. H. Berry. 4, UZMK (unregistered),

Ceylon and Calcutta, lat. 20°51'N, long. 87°58'E, coll. Th.

Mortensen, 26.7.51. 2, UZMK (unregistered), Ceylon and Calcutta, lat. 20°51'N, long. 87°58'E, coll. Galathea Expd.,

26.4.65. 3, UZMK (unregistered), Bombay, lat. 18°48'N, long.

72°37'E, 20-30 m, coll. Galathea Expd., 12.5.64. 1, ANSP

77146, Bombay, coll. F. F. Hallberg. 1, ANSP 77546, Bombay,

coll. F. F. Hallberg. 1, ANSP 77520, Bombay, coll. F. Hallberg. 1, ANSP 88356, Bombay, coll. H. W. Fowler.

PAKISTAN: 1, BMNH 1882:2.1.4060, Sind, coll. F. Day.

PERSIAN GULF: 10, BMNH 1911.2.23.58-67, Persian Gulf,

coll. Tounsend. 2, BMNH 1928.3.20.89-90, N end of Persian

Guf, ex. Indian Museum.

27. Cynoglossus robustus Gunther

FIGURE 30; PLATE 12

Cynoglossus robustus Gunther, 1873a:243 [type-locality:

Shanghai].—Steindachner, 1896:219 [Japan].—Jordan and

Snyder, 1901:123.—Jordan and Starks, 1906a:239 (Japan,

China].—Fowler and Bean, 1922:68 [Tokyo]. Jordan and

Hubbs, 1925:302 [Kobe].—Fowler, 1930:616 [Hong Kong].—

Wu, 1931:169 [Chi-Pou].—Chu, 1931:95 [Shanghai, Hong

Kong].—Wu, 1932:147 [Tehou-San].—Wu and Wang,

1933:304 [Amoy].—Fowler, 1934b:221 [China].—Okada and

Matsubara, 1938:437 [Chinese sea].—Mori, 1952:183 [Quelpar Is.].—Kamohara, 1952:86 [Tosa].—Matsubara, 1955:1285

[S Japan. S China Sea].—Tchang, 1955:297, fig. 183.—Chen,

1956:106 [Kaohsiung].—Ochiai, 1959:213.—Chu, 1963:539,

fig. 405.—Chen and Weng, 1965:96, fig. 67 [Kulung].—

Shen, 1967:213, figs. 130-133 [Kowloon].

Cynoglossus brunneus Regan, 1905a:26 [Inland Sea of Japan].

—Jordan and Starks, 1906a:239 [Inland Sea of Japan].—

Jordan, Tanaka and Snyder, 1913:336 [listed].

Cynoglossus inusita Jordan, Tanaka, and Snyder, 1913:335

[type-locality: Tokyo, Nagasaki].—Hubbs, 1915:494 [Kobe].

DESCRIPTION.—Based on 16 specimens, 100.0-388.0

mm SL, including the holotype of C. robustus.

Depth of body 21.03-29.25 (M = 25.09), length

of head 20.16-24.46 (M = 22.46) percent of standard

length. Diameter of eye 6.52-9.88 (M = 7.84), interorbital width 3.70-7.64 (M = 5.37) percent of length

of head. Anterior nostril tubular, in front of lower

eye, posterior nostril simple, in the anterior half

of the interorbital space. Snout obtusely pointed

33.58-44.83 (M = 39.36) percent of length of head,

rostral hook short, extending only to front of

anterior nostril. Maxillary extending to beyond

fixed eye; angle of mouth extending to just vertical

from below posterior border of fixed eye, nearer

to branchial opening than to tip of snout; tip of

snout to angle of mouth 48.91-56.32 (M = 53.64)

angle of mouth to branchial opening 45.00-57.94

(M = 47.99) percent of length of head.

Scales: Ctenoid on ocular side with the lateral-line

scales cycloid anteriorly, ctenoid posteriorly; scales

on blind side cycloid.

Lateral-Line System: Two lateral lines on eyed

side, midlateral line with 70-82 scales, 10 scales

between. No lateral line on blind side.

lem

FIGURE 30.—Outline drawing of C. robustus, holotype (BMNH 1873.730.61)

from Shanghai, China.
NUMBER 238 65

Interlinear scale rows 10

Frequencies 8

Fins: Dorsal with 125-134 (M = 129) rays, anal

with 90-104 (M = 100) rays, caudal 10 in 8 specimens (radiographs).

Vertebrae: 56-61 comprising 10 (rarely 9) abdominal and 47-51 caudal elements in 8 specimens

(radiographs).

Coloration: Upper side light brown, lower whitish

in preserved specimens.

Size: Largest specimen examined, 413 mm, is from

Onomichi, Japan.

DISTRIBUTION.—From South China Sea through

Taiwan, Tung-hai, Yellow Sea, Po-hai to Korea

and Japan.

DIAGNOSIS AND AFFINITIES.—Cynoglossus robustus

is closely allied to C. arel. It differs from C. arel in

having a more elongate body, longer snout, and

smaller eyes with a wider interorbital space (3.70-

7.94, M = 5.37 percent of length of head cf. 1.75—

5.21, M = 3.43 percent). The number of interlinear

scale rows further distinguishes the two species, C.

robustus having an invariable number of 10, while

C. arel has a count of 7-9 (M = 7).

NOTE ON SYNONYMY.—Gunther's (1873a:243) C.

robustus was described on the basis of a specimen,

212.5 mm in TL, from Shanghai and characterized

as having two lateral lines on ocular side, separated

by 10 longitudinal series of scales, and one lateral

line on blind side. From the Inland Sea of Japan,

Regan (1905a:26) described C. brunneus based on

a single specimen, 200 mm in TL. The salient

feature in which this fish is differentiated from C.

robustus is its slightly larger scales, with only 9

interlinear scale rows. I have examined the holotype

of C. brunneus and found it to possess 10 scale rows

between the middle and the upper lateral lines;

therefore there can be no doubt that they are

identical. Jordan and Starks (1906a: 239) recorded

C. robustus both from Japanese and Chinese waters

but doubted the conspecificity of the Chinese C.

robustus with that of the Japanese. Jordan, Tanaka,

and Snyder (1913), however, gave a new name, C.

inusita, to the Japanese specimens procured by

Jordan and Starks. Hubbs (1915:494) concurred

with Jordan and collaborators in keeping the

Japanese form distinct and further pointed out

that the Japanese specimens differed from the

type specimens of C. robustus in having fewer rays

(122-127) and scales (74) on the midlateral line,

whereas the type specimens have 130 rays in the

dorsal and 83 scales along the midlateral line. The

differences in the fin rays and scale count noted by

Hubbs in the Japanese specimens in comparison

with the type specimens of C. robustus is not, however, substantiated by the more numerous specimens

now available.

TYPE SPECIMENS.—Holotype of C. robustus,

BMNH 1873.7.30.61, 296.5 mm SL, from Shanghai,

China, coll. Swinhoe. Holotype of C. brunneus,

BMNH 1905.6.6.248, 186 mm SL, from Inland Sea

of Japan, coll. Gordon Smith.

OTHER MATERIAL EXAMINED.—CHINA: 1 specimen, BMNH

1862.11.1271, Shanghai, coll. Jamrach. 1, BMNH 1925.526.17,

Yenting, China, ex. Sci. Soc, China. 1, BMNH 1031.12.1268,

Chusen Is., China, coll. S. L. Hora (Indian Museum). 1,

USNM 27016, China, coll. Jordan. 3, USNM 159452, Yellow

Sea, China, coll. C. L. Hubbs, 1929. 1, KU 20726, E China

Sea, ex. Kyoto University. 1, MCZ 11329, Hong Kong,

coll. Capt. Putnam. TAIWAN: 1, USNM 7660, Taiwan, coll.

F. Beker. 1, USNM 177429, Taipei Mkt., Taiwan, coll.

D. K. Lawless. KOREA: 1, UMMZ 159452, Korea, near

Fusan, coll. C. L. Hubbs, 1929. 4, UMMZ 159450, Korea,

coll. C. L. Hubbs, 1929. JAPAN: 1, BMNH 1923:2.26.649,

Tokyo, Japan, coll. Jordan. 1, USNM 37886, Tomo, Japan.

1, USNM 39966. Japan, coll. Hitchcock. 2, USNM 56386,

Japan, coll. Jordan and Snyder. 2, USNM 56349, Onomichi,

Japan, coll. Jordan and Snyder. 1, USNM 75933, Japan. 1,

USNM 77145. Kobe, Japan, coll. Albatross, 1906. 1, USNM

159449, Osaka Mkt. Japan, coll. C. L. Hubbs, 7.10.1929. 1.

USNM 159453, Kobe Mkt. Japan, coll. C. L. Hubbs, 1929.

2, USNM 159690, Mikawa Bay, Japan, coll. M. Tshikawa,

1922. 1, KU 12612, Kofu, Japan, coll. Kyoto University,

Jan 1950.

28. Cynoglossus lingua Hamilton-Buchanan

FIGURE 31; PLATE 12

"Jerree Potoo."—Russell, 1803:57, pi. 73.

Cynoglossus lingua Hamilton-Buchanan, 1822:32, 365 [typelocality: Gangetic estuaries].—Vinciguerra, 1889:190.—

Gunther, 1862:501.—Day, 1877:433, pi. 96: fig. 1; 1889:454,

fig. 163.—Norman, 1928:200, fig. 16.—Weber and de Beaufort, 1929:203.—Smith, 1933:83.—Suvatti, 1936:97; 1950:327.

—Munro, 1955:264.—Fowler, 1956:186—Scott, 1959:42.—

Kuronuma, 1961:32.—Pradhan, 1964:458 [Bombay coast].—

Punpoka, 1964:62, fig. 20.

Plagusia lingua.—Cantor, 1850:1215.—Jerdon, 1853:148.

Arelia lingua.—Kaup, 1858:107.

Pleuronectes potous Cuvier, 1836:307.

Plagusia potous.—Bleeker, 1852:23.—Jerdon, 1853:148.

Arelia potous.—Sleeker, 1859:185.

Cynoglossus potous.—Bleeker, 1875:33, pi. 241: fig. 4.

Plagusia macrorhynchos Bleeker, 1851a:413 [type-locality:

Tjilankahan]; 1852:22.
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 31.—Outline drawing of C. lingua (BMNH 1933.7.31.25) from Singapore.

Arelia macrorhynchos.—Kna, 1867:295.—Bleeker, 1875:3, pi.

242: fig. 2.—Weber and de Beaufort, 1929:208.

Cynoglossus acinaces Jenkins, 1910b:130 [type-locality: Khulna,

£. Bengal].

DESCRIPTION.—Based on 18 specimens, 214.0-345.0

mm SL.

Depth of body 17.24-21.71 (M = 18.94), length of

head 21.17-25.75 (M = 22.95) percent of standard

length. Diameter of eye 5.26-8.51 (M • 6.51), interorbital width 1.75-6.40 (M = 3.28) percent in length

of head. Anterior nostril of eyed side tubular, in

front of lower eye, on upper lip, posterior nostril

simple, in anterior half of interorbital space. Snout

pointed, 37.04-45.00 (M = 41.21) percent of length

of head, rostral hook rather short, extending just

below the anterior nostril. Maxillary extending

well beyond fixed eye; angle of mouth extending to

just beyond vertical from posterior border of fixed

eye, nearer to branchial opening than to tip of

snout; tip of snout to angle of mouth 50.00-58.33

(M = 53.81), angle of mouth to branchial opening

40.47-49.09 (M = 45.57) percent of length of head.

Scales: Ctenoid on ocular side with cycloid scales

on head and on the lateral lines; cycloid on blind

side.

Lateral-Line System: Two lateral lines on ocular

side, midlateral line with 90-101 scales, 11-12

scales between them. No lateral line on blind side.

The midlateral line on the ocular side, instead of

opening through simple pores on every scale, opens

by means of ducts, one on either side into the

adjoining scale (Figure 2).

Interlinear scale rows 11 12

Frequencies 5 16

Fins: Dorsal with 126-138 (M = 132) rays, anal

with 97-114 (M = 101) rays, caudal 10 in 17 specimens (radiographs).

Vertebrae: 57-66, comprising 9 abdominal and

48-57 caudal elements in 17 specimens (radiographs).

Coloration: Upper side uniformly brown with or

without darker patches, lower lighter in preserved

specimens.

Size: Largest specimen examined, 378 (355 + 23)

mm, is from Indonesia.

DISTRIBUTION.—From Malay Archipelago including Thailand and Vietnam to seas and estuaries

of India and Pakistan.

DIAGNOSIS AND AFFINITIES.—Cynoglossus lingua

is closely related to the other two species of the

arel complex, but it is immediately distinguished

from them by the marked elongation of the body,

a longer snout, much smaller eyes, and smaller

body scales having an interlinear scale count of

11-13<M = 12).

NOTE ON SYNONYMY.—Hamilton-Buchanan (1822:

32) described C. lingua from the Gangetic estuaries,

based on his unpublished original drawings now

preserved in the Asiatic Society of Bengal and

reproduced by Hora (1929, pi. 15: figs. 1, 2). Norman

(1928:200) included in the synonymy Pleuronectes

potous Cuvier, Plagusia macrorhynchos Bleeker,

C. elongatus Giinther, and C. acinaces Jenkins. I

concur with Norman in including all these species

as synonyms of C. lingua except C. elongatus, which

I have synonymized with C. arel.

Bleeker (1851a:4l3) described P. macrorhynchos

on the basis of a figure of the fish from Tjilankakan,

Bantam. There is no specimen of the fish in Leiden

(pers. comm., Dr. M. Boeseman). Cuvier's description of P. potous was based on Russel's "Jerree
NUMBER 238 67

Potoo" and not on any specimen. Jenkins (1910b:

130) described C. acinaces on the basis of five

specimens, 59-148 mm SL, from Morrelganj, Kulna

district, Bangladesh and related it to C. elongatus

and C. lingua but differentiated them in the much

higher number of rays in the dorsal and anal fins

and in the greater slenderness of the body. The

differences noted by Jenkins fall within the normal

range of variation of C. lingua.

TYPE SPECIMENS.—Gunther (1862:501) considered a stuffed specimen from the Ganges in the

British Museum, presented by G. R. Whitehouse,

as probably the type of C. lingua. Gunther (1861:

IV), referring to the Whitehouse collections, was

very cautious saying that he "believed [it] to contain

many typical specimens of Hamilton-Buchanan's

work." Hora (1929) examined this question and

concluded that there was absolutely no evidence

to show that the Whitehouse collections contained Hamilton-Buchanan's specimens. HamiltonBuchanan never kept any zoological collections

and for his descriptions in the "Gangetic Fishes"

he relied entirely on his drawings and field notes.

I have not been able to locate the stuffed specimen

referred to by Gunther.

A specimen ZSI F 4149/1, 148 mm SL, the largest

of the syntypes, from Sunderbans of Kulna district,

Bangladesh, is selected here as the lectotype of C.

acinaces. The other four specimens, 59-114 mm SL,

are paralectotypes.

OTHER MATERIAL EXAMINED.—CHINA: 1 specimen, BMNH

1862.11.1.271, Shanapai, coll. Jamrack. INDONESIA: 2.

USNM 72748, Batavia, Java, coll. O. Brayant, 4.2.1909. 1,

ZMA 108005, Indonesia, coll. P. Bleeker, 1806. 1, ZMA

108006, Indonesia, coll. Gier Expd.. 1908. 2, MCZ 30812,

Batavia, coll. Brayant and Palmix. BORNEO: 2, USNM

137410 and 13411, Luzon and Borneo, coll. Albatross. THAILAND: 3, USNM 103320, Thailand, coll. H. M. Smith.

23.6.1933. 2, USNM 119682, Thailand, coll. H. M. Smith,

23.6.1933. 5, ANSP 76788, Thailand, coll. R. M. Schauensee,

1936. 1, ANSP 76813, Thailand, coll. R. M. Schauensee,

1936. 3, ANSP 87318, Thailand, coll. R. M. Schauensee, 1936. 2, ANSP 274537, Bangkok, coll. R. M. Schauensee.

6, ANSP 89447, Bangkok, coll. R. M. Schauensee and H. W.

Fowler, 1936. 5, CAS (GVF), Gulf of Thailand, coll. local

fishermen, 7.9.1957. 3, CAS (GVF) 2582, lat. 13°26'N. long.

lOClO'E, coll, Naga Expd., 17.5.61. 1, MCZ 11338, Thailand,

coll. C. L. Salmin, 1871. 1, UMMZ 181230, Gulf of Thailand,

coll. John Bardach. 1, BMNH 1934.12.18.78, Gulf of

Siam, coll. Siamese Museum. BURMA: 3, MCZ 11134, Rangoon, coll. Theobals. 1, BMNH 1870.6.14.7, Moulmein, coll.

F. Day. 1, ZSI

 F l511

, off Akyab, 34 fms, coll. Marine Survey. SINGAPORE: 1, MCZ 11513, Singapore, coll. Capt.

Putnam. 3, BMNH 1933.7.3124-26, Singapore Mkt.. coll.

Raffles Museum. VIETNAM: 1. TNSP 76858. Saigon,

coll. Hugo Rutherford, Dec 1934. MALAYA: 4. MCZ 274867.

Penang, coll. Capt. Putnam. 1, BMNH 1938.8.10.1, Penang,

coll. Fish Inst. 1, BSI 9055, Penang, coll. F. Stoliczka.

INDIA: I, BMNH 1855.12.26.601. Ganges, coll. Zool. Soc. 1,

BMNH 1872.4.17.37, N. E. Bengal, coll. Jordan. 2, BMNH

1889.2.1.4051, 4055, Calcutta, coll. F. Day. 1, BMNH

18892.1.4054. Calcutta, coll. F. Day. 1, BMNH 1889.2.1.4056.

Orissa, coll. F. Day. 1, BMNH 1889.2.1.4057, Madras, coll.

F. Day. 2. BMNH 1928.3.20.85-86, locality not given, ex.

Indian Museum. 1, ZSI 1461, Calcutta, coll. F. Day. 1,

T* 1 fil ft

ZSI —2— , Mouth of Arasalar R., Karaikal. coll, A. G. K.

Menon, 10.3.58. l.ZSI

 F ^6 5 °. Sunderbans. coll. A. Rahim.

1, ZSI—-—, Sullermukai R., 24-Pargans, coll. J. T. Jenkins.

F I7Q1

16. ZSI , Karar, N Kanara Dt., Maharastra, coll. K. K.

F 1649

Tiwari, 20.2.56 6, ZSI . Sunderhans. coll. S. W. Kemp.

l.ZSI —— , Pulta Water Works, Barrackpore. coll. Pulta

F 1040

Survey. l.ZSI ^-y^ , 24-Parganas. coll. Bengal Fisheries. 1.

ZSI

 F 5849

, 24-Parganas, coll. J. T. Jenkins. 1, ZSI

 F 5987

,

Nabaspur, coll. J. T. Jenkins. 1, specimen, ZSI (unregistered). Morrison Bay. coll. Chilka Survey. 1, ZSI 1208,

Calcutta.

The cynoglossus group

This is a group of specialized species of Cynoglossus characterized by relatively small eyes (pedunculate in monopus) separated by a small interorbital

space, a moderately elongate snout with the angle

of mouth situated nearer tip of snout than to

branchial opening (except lida), two lateral lines

on ocular side, and relatively small ctenoid scales

on both sides of the body. No lateral line on blind

side.

Included in this group are the cynoglossus,

monopus, puncticeps, and lida complexes. Their

combined range covers the whole of the Malay

Archipelago, the seas of India and Pakistan, and the

East Coast of Africa. This group appears to be

related to the axel complex and may have evolved

from a recent offshoot of the axel stock.

The cynoglossus complex

Cynoglossus cynoglossus, C. semifasciatus, and C.

maexostomus form the cynoglossus complex. The

characteristics of this complex are the small eyes
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

separated by a small interorbital space, obtusely

pointed snout with the angle of the mouth nearer

to the tip of snout than to the branchial opening,

two lateral lines on ocular side, and relatively small

scales on body, the interlinear scale count being 12

to 16. This complex ranges through the Malay

Archipelago and the seas of India and Pakistan.

29. Cynoglossus cynoglossus (Hamilton-Buchanan)

FIGURE 32; PLATE IS

Achirus cynoglossus Hamilton-Buchanan, 1822:132, 373 [typelocality: Ganges mouth].

Plagusia cynoglossus.—Cantor, 1850:1211.

Icania cynoglossa.—Kaup, 1858:109.

Cynoglossus cynoglossus.—Norman, 1928:208 [Bengal].—

Weber and de Beaufort, 1929:199.—Punpoka, 1964:60.

Plagusia oxyrhynchos Bleeker, 1851a:416 [type-locality:

Batavia]; 1852:26.

Arelia oxyrhynchos.—Bleeker, 1859:185.

Cynoglossus oxyrhynchos.—Gunther, 1862:499.—Bleeker,

1875:36, pi. 245: fig. 1.—Weber and de Beaufort, 1929:201.

Plagusia sumatrana Bleeker, 1853c:529 [type-locality: Benculen, Sumatra].

Arelia sumatrensis.—Bleeker, 1859:185.

Cynoglossus sumatrensis.—Gunther, 1862:497.—Bleeker,

1875:35, pi. 243: fig. 1.—Jordan and Richardson, 1908:281

[Ticao Island].—Fowler, I918b:65 [Philippines].

Cynoglossus sumatranus.—Norman, 1928:209.—Weber and de

Beaufort, 1929:202.—Fowler, 1938:88.—Herre, 1953:191.—

Punpoka, 1964:72.

Plagusia bengalensis Bleeker, 1853a:152 [type-locality:

Hooghly, Calcutta].

Cynoglossus bengalensis.—Gunther, 1862:499 [Ganges].—Day,

1877:435, pi. 97: fig. 4 [Hooghly, Calcutta].

Cynoglossus hamiltonii Gunther. 1862:504 [Ganges:

Pinang].—Duncker, 1904:169.

Cynoglossus buchanani Day, 1869:522 [India].

Cynoglossus deltae Jenkins, 1910b: 130 [type-locality: Khulna,

E Bengal].—Norman, 1928:212.

Cynoglossus semifasciatus [not Day].—Suvatti, 1936:98; 1950:

328.—Punpoka, 1964:70.

DESCRIPTION.—Based on 45 specimens, 62.0-

147.0 mm SL, including the lectotype and paralectotype of C. deltae.

Depth of body 22.03-34.21 (M = 27.11), length of

head 18.94-28.95 (M = 22.54) percent of standard

length. Diameter of eye 3.03-13.04 (M = 8.44), interorbital space 1.89-9.68 (M = 3.56) percent of

length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior nostril simple, either in front of interorbital

space or in anterior half of it. Snout obtusely

pointed, 21.43-45.45 (M = 32.87) percent of length

of head, rostral hook rather short, extending to

front of anterior nostril or reaches just short of

perpendicular through anterior border of fixed

eye. Maxillary extending to below posterior border

of fixed eye or just behind it; angle of mouth

extending below vertical from posterior half of

fixed eye, nearer to tip of snout than to branchial

opening, tip of snout to angle of mouth 39.22-

53.49 (M = 45.64), angle of mouth to branchial

opening 47.37-60.71 (M = 51.97) percent of length

of head.

Scales: Ctenoid on both sides.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line usually undulating and stopping at a very short distance from caudal base,

midlateral line with 70-90 (M = 82) scales, 12-14

(M = 13) scales between them. No lateral line on

blind side.

FIGURE 32.—Outline drawing of C. cynoglossus (BMNH 1858.8.15.55)

from Ganges, coll. G. K. Waterhouse.
NUMBER 238 69

Interlinear scale rows

Frequencies

12 13 14

6 8 26

Fins: Dorsal with 95-102 (M=100) rays, anal

with 72-78 (M = 75) rays, caudal 10 in 42 specimens

(radiographs).

Vertebrae: 44-48, comprising 9 abdominal and

35-39 caudal elements in 42 specimens (radiographs).

Coloration: Upper side uniformly brownish with

or without darker spots, lower whitish in preserved

specimens.

Size: Largest specimen examined, 160 (147 + 13)

mm, is from the Ganges.

DISTRIBUTION.—From Malay Archipelago to Philippines, and westward to Burma, Bangladesh, and

India (W Bengal). In seas and estuaries as far as

the tidal reaches.

DIAGNOSIS AND AFFINITIES.—The nearest relative

of C. cynoglossus is C. semifasciatus. The species,

however, differs in several respects, especially in

the obtusely pointed and longer snout (mean

length 32.87 percent of length of head cf. 27.40

percent), wider interorbital space (mean width

3.56 percent of head cf. 2.16 percent), and the

greater number of interlinear scale rows (12-14,

M = 13 cf. 11-14, M = 12).

NOTE ON SYNONYMY.—Hamilton-Buchanan (1822:

132) described C. cynoglossus from the Ganges on

the basis of his original drawing of the fish now

in the Asiatic Society of Bengal and reproduced by

Hora (1929, pi. 19: figs. 2, 3). Norman (1928:208)

synonymized Plagusia bengalensis Bleeker with C.

cynoglossus and Bleeker's P. oxyrhynchos and P.

sumatrana are here included under C. cynoglossus.

Bleeker (1851a:416) characterized P. oxyrhynchos

by its two lateral lines on the left side separated by

13 longitudinal series of scales, a single lateral line

on the blind side, two nostrils, and subcontiguous

eyes; and P. sumatrana (Bleeker, 1853c:529) with

two lateral lines on the left side, one on the right,

two nostrils, and a wider interorbital space. He,

did not, however, indicate the interlinear scale rows

in P. sumatrana. I have examined a specimen of P.

sumatrana (BMNH 1862.6.3.9) and another of P.

oxyrhynchos (BMNH 1862.6.3.17) from the Bleeker

collection in the British Museum and compared

them with the specimens of C. cynoglossus from the

Ganges, and I have no hesitation in synonymizing

them. In the specimens examined, the lateral line

on the blind side, stated to be present on Bleeker's

P. oxyrhynchos and P. sumatrana, is absent and in

all other respects they conform well with C. cynoglossus.

Jenkins (1910b: 130) C. deltae was described on

the basis of two specimens from Morrelganj, Kulna,

Bangladesh, and he characterized it by two lateral

lines on the ocular side separated by 10-12 series

of scales. In the interlinear count, as well as in all

other meristic and proportional measurements, the

types of C. deltae agree so well with C. cynoglossus

that they are undoubtedly conspecific.

Gunther's (1862:504) C. hamiltonii, described on

the basis of a dried skin from Cantor's collection

from Penang in the British Museum, and characterized with two lateral lines on the left separated

by 13 series of scales is synonymized here with C.

cynoglossus. Giinther has not noted any conspicuous

nostril in his specimen. I have not been able to

trace the type (skin) mentioned in Gunther's catalog to ascertain whether the nostril is present or

not. From the other details of the description, however, I have come to a definite conclusion that C.

hamiltonii is nothing other than C. cynoglossus.

TYPE SPECIMENS.—As stated earlier, HamiltonBuchanan never kept any zoological collections and

for his descriptions he entirely relied on his drawings and field notes. Cynoglossus cynoglossus was

not figured in his book on the fishes of the Ganges

(1822) either. Hora (1929:pl. 19: figs. 2 and 3),

however, reproduced Hamilton-Buchanan's manuscript drawing of C. cynoglossus, a specimen measuring about 106 mm long.

Bleeker's description of P. oxyrhynchos was based

on five specimens, 100-120 mm in TL from Batavia.

Bleeker in his Atlas (6:36, pi. 245: fig. 1) redescribed the species on eight specimens, 100-120 mm

in TL from Batavia, Borneo, and Amboina and illustrated it.

In the Bleeker collection in Leiden there are six

specimens, 88-111 mm SL, 97-123 TL, of P. oxyrhynchos cataloged as RMNH 6793. Chabanaud in

1946 indicated a specimen, 92 mm SL, 101 mm TL,

as the lectotype. The largest specimen estimated

here as measuring 123 mm TL may have been the

"120" specimens that Chabanaud seems to have rejected because of damage. The 92 mm SL specimen

is recognized as the lectotype and retained as

RMNH 6793, the remaining five specimens including the "120" example having been removed to

RMNH 26210. Of these five, two, 58-90 mm S.L.,
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

97-99 mm TL, are too small to be considered as

paralectotypes; the other three specimens, 93-111

mm SL, 103-123 mm TL, are recognized as paralectotypes. (Dr. M. Boeseman examined the type

material on my behalf.)

Bleeker (1853c:526) described P. sumatrana based

on a single specimen 121 mm long from Benculen,

Sumatra. In the Bleeker collection in Leiden there

is a specimen cataloged as RMNH 6787, which unfortunately is mutilated with the head mostly

missing, but otherwise agrees with Bleeker's figure

of a 121 mm example. Strangely Bleeker, in the

Atlas (6:36), refers to only a single specimen though

he lists three localities. According to Dr. Boeseman,

at the 1879 auction there was only a single specimen in the Bleeker collection and since it agrees

with Bleeker's figure in size, it must be the holotype.

Bleeker's description of P. bengalensis was based

on two specimens, 120 mm TL and 139 mm TL,

from Hooghly River near Calcutta. According to

Dr. Boeseman, in the Bleeker collection in Leiden

there are two examples, 106 mm SL, 120 mm TL

and 126 mm SL, 139 mm TL, both syntypes, cataloged as RMNH 6794. The largest specimen is selected here as the lectotype, and is retained as

RMNH 6794; the other specimen recataloged as

RMNH 26211 is the paralectotype.

A specimen, 64.0 mm SL, ZSI F 4150/2 from

the Sunderbans of Morelganj, Khulna district,

Bangladesh, collected by Bengal Fisheries, is selected as the lectotype of C. deltai and a specimen,

BMNH 1928.3.20.133, 62.5 mm SL, taken along

with the lectotype as the paralectotype.

OTHER MATERIAL EXAMINED.—PHILIPPINES: 1 specimen,

ANSP 49038, Philippines, coll. R. M. de Schauensee. 1,

NHV 43826, Philippines, coll. Naga Expd. INDONESIA: 1,

BMNH 1862.6.3.9, Sumatra, coll. P. Bleeker. 1, BMNH

1862.6.3.17, locality not given, coll. P. Bleeker. 1, BMNH

1931.4.23.46, Batavia Mkt., coll. Hardenberg. 1, ZMA

108.007, Indonesia, coll. Kleiweg de Zwaan. 1, AMNH 18414,

Java, coll. Zeeviss. 1, USNM 87943, Sumatra, coll. H. C.

Rollers. THAILAND. 2. ANSP 59918, Thailand, coll.

R. M. de Schauensee. 3, ANSP 61843-51, Thailand, coll.

R. M. de Schauensee. 1, ANSP 87278, Thailand, coll. R. M.

de Schauensee. 4, ANSP 87840. Thailand, coll. R. M. de

Schauensee. 31, ANSP 89461, Thailand, coll. R. M. de

Schauensee. 6, CAS (GVF), Gulf of Thailand, coll. Naga

Expd. 1, CAS (GVF), Thailand, coll. Naga Expd. 1, NHV

43766, Thailand, coll. Naga Expd. MALAYA: 2, MCZ

30343, Penang coll. E. P. Ward. BURMA: 1, BMNH

1928.3.20.112, Mergui, coll. Indian Museum. 2, ANSP 77027.

Burma, coll. Halberg. 4, MCZ 33199, Burma. INDIA: 1,

BMNH 1852.9.13.235A, Hooghly R., coll. Zool. Society. 1,

BMNH 1855.9.19.12.32. Ganges, coll. Hasler. 1, BMNH

1855.12.26.602, R. Ganges, coll. Zool. Soc. 1, BMNH

1862.6.3.9. Sumatra, coll. P. Bleeker. 1, BMNH 1862.6.3.17,

locality not given, coll. P. Bleeker. 2. BMNH 1889.2.1.4065-

66. Calcutta, coll. F. Day. 1. BMNH 1928.3.20.133.

Sunderbans, ex. Indian Museum. 2, BMNH 1928.3.20.108

and 112, Calcutta, ex. Indian Museum. 3, BMNH 1928.3.20.

109-111, Hooghly R., ex. Indian Museum. 1, BMNH

1934.11.21.8c, Hooghly R., ex. Zoological Survey of India. 2.

BMNH 1954.5.20.21, Hooghly R., ex. Zoological Survey of

India. 1, ZSI 461. Calcutta, coll. J. Anderson. 1. ZSI 1499,

Calcutta, coll. F. Day. 1, ZSI 2689, Calcutta, coll. F. Day. 2,

ZSI 752/2, Hooghly R., coll. Hooghly Survey. 2. ZSI

(unregistered), Tolly's Nallah, coll. Nirod. 1. ZSI F 2014/2,

Barrackpore. coll. Pulta Survey. 2. ZSI F 3434/1, Orissa

coast, coll. C. A. Paiva. 6, ZSI F 5042/2, Sunderbans,

Gangetic delta, coll. S. W. Kemp. 1, ZSI 6865/2, Sunderbans,

Gangetic delta, coll. S. W. Kemp.

30. Cynoglossus semifasciatus Day

FIGURE S3; PLATE 13

Cynoglossus semifasciatus Day, 1877:436, pi. 97: fig. 5 [typelocality: 'Sea at Madras']; 1889:458.—Alcock, 1889:289.—

Jenkins, 1910a:30.—Norman, 1928:207.

Cynoglossus brevirostris Day, 1877:437, pi. 97: fig. 6 [typelocality: Madras]; 1889:459.—Norman, 1928:212.

Cynoglossus bengalensis [not Bleeker].—Johnstone, 1904:209.

Cynoglossus cynoglossus [not Hamilton-Buchanan].—Munro,

1955:266. pi. 50 [Ceylon].—De Silva. 1956:197 [Ceylon].

DESCRIPTION.—Based on 41 specimens, 75.5-136.0

mm SL, including the holotype of C. semifasciatus

and C. brevirostris.

Depth of body 24.53-31.58 (M = 27.88), length of

head 19.81-29.41 (M = 22.76) percent of standard

length. Diameter of eye 5.36-12.96 (M = 8.29), interorbital space narrow, 1.79-2.50 (M = 2.21) percent of length of head. Two nostrils on ocular side,

anterior nostril tubular, in front of lower eye, posterior nostril simple, in the anterior half of interorbital space. Snout rounded or obtusely pointed,

21.43-32.50 (M = 27.40) percent of length of head,

rostral hook rather short, extending to front of

anterior nostril. Maxillary extending well beyond

posterior border of fixed eye; angle of mouth extending a little beyond vertical from fixed eye,

nearer to tip of snout than to branchial opening;

tip of snout to angle of mouth 33.33-47.62 (M

= 41.66), angle of mouth to branchial opening

47.62-60.73 (M = 54.64) percent of length of head.

Scales: Ctenoid on both sides, including those of

lateral lines.
NUMBER 238 71

FIGURE 33.—Outline drawing of C. semifasciatus (BMNH 19282.6.1) from Tanur, Malabar coast.

Lateral-Line System.—Two lateral lines on ocular

side, the dorsolateral line entering the dorsal fin

at a short distance from the caudal base, the midlateral line with 70-78 (M = 75) scales, 11-14 (M

= 12) scales between them in 31 examples. No lateral line on blind side.

Interlinear scale rows 11 12 13 14

Frequencies 2 16 11 2

Fins: Dorsal with 99-107 (M = 102) rays, anal

with 75-83 (M = 79), caudal 10 in 23 specimens

(radiographs).

Vertebrae: 47-50 comprising 9 abdominal and

38-41 caudal elements in 23 specimens (radiographs).

Coloration: Upper side light brownish with a

number of irregular vertical dark bands, lower

whitish in preserved specimens.

Size: The largest specimen examined from Portonovo, Madras, trawled by Anton Bruun is 138

(123+15) mm long.

DISTRIBUTION: East coast of India and Ceylon,

recorded from a depth of 7-10 fms.

DIAGNOSIS AND AFFINITIES.—In many morphometric characters C. semifasciatus resembles C. macrostomus. It is distinguishable, however, by its deeper

body (mean depth 27.88 percent of SL cf. 25.89

percent), lesser number of interlinear scales (11-14,

M = 12 cf. 14-16, M = 15), and midlateral scales

(70-78, M = 75 cf. 80-92, M = 83).

NOTE ON SYNONYMY.—Day (1877:436) described

C. semifasciatus on the basis of a single specimen

from Madras coast and characterized it as having

two lateral lines on ocular side separated at then

highest distance by 12 or 13 rows of scales. Norman

(1928), in his revision of flatfishes of India, noted

that in C. semifasciatus the lateral lines on the

ocular side are separated by 13-14 series of scales

and not 12 or 13 as Day observed. The name C.

semifasciatus, however, had been wrongly assigned

to the commercial species of the Malabar coast,

namely, C. macrostomus, having wider range of

15-16 scales between the lateral lines (Menon 1971).

This was mainly brought about by Norman's (1928:

207) placing C. semifasciatus in the group, showing

a range of 12 or 15 scales between the lateral lines

in his key to the species of Cynoglossus; thereby,

workers using Norman's key, having assumed a

wider range for C. semifasciatus, confused the

Malabar sole for this species. Cynoglossus semifasciatus can be readily distinguished by its lower

number of interlinear scales, 11-14 (M = 12) cf.

14-16 (M = 15).

Day (1877:437) described another species from

Madras coast, C. brevirostris, based on a single

specimen and characterized it as having a lesser

number of 10 scale rows between the lateral lines

on ocular side. I have examined the holotype of

C. brevirostris and found it to have 11 interlinear

scales, which falls within the range of variation of

this character for C. semifasciatus. I have, therefore,

included C. brevirostris in the synonymy of C.

semifasciatus.

TYPE SPECIMENS.—Holotype of C. semifasciatus,

ZSI 2490, 102 mm SL, Madras coast, coll. F. Day.
72

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Holotype of C. brevirostris, ZSI 2690, 110 mm SL,

Madras coast, coll. F. Day.

OTHER MATERIAL EXAMINED.—INDIA: 2 specimens, ZSI

2359/2, 2360/2, Marina Beach, Madras, coll. ZSI, Southern

Regional Sta. 1, ZSI F 3437/1, Pun, Orissa coast, coll.

Golden Crown. 2, ZSI 12680/2, Gopalpur, Orissa, coll.

Investigator. 2, ZSI (unregistered), Madras, coll. A. Daniel.

28, ZSI (unregistered), Waltair coast, coll. K. V. Sekharan.

2, BMNH 1889.2.1.4066-67, Calcutta, coll. F. Day. 1, BMNH

1928.3.20.107, Ganjam coast, coll. Investigator. 1, SOSC,

Ref. 334, Thirumulli, Madras, lat. 11°13'N, long. 79°53'E,

coll. Anton Bruun, 23.9.1966. 3, SOSC Ref 334, Madras,

lat. 11°35'N, long. 79°5'E, coll. Anton Bruun. 1, SOSC

Ref 334, Vellar Estuary Madras, lat. ll°30'N, long 79°50'E,

coll. Anton Bruun. 4, SOSC Ref 334, Porto Novo, Madras,

coll. Anton Brunn.

31. Cynoglossus macrostomus Norman

FIGURE 34; PLATE 14

Cynoglossus hamiltonii [not Gunther].—Day, 1877:436, pi. 95:

fig. 3 ["Hooghly at Calcutta"]; 1889:458.

Cynoglossus macrostomus Norman, 1928:204 [type-locality:

Hooghly Estuary, near Calcutta].

Cynoglossus luctuosus Chabanaud, 1947e:813 [type-locality:

Madras].

Cynoglossus semifasciatus [not Day].—Seshappa and Bhimachar, 1955:183.—Saramma, 1963:77 [Kerala coast].

Cynoglossus cynoglossus [not Hamilton-Buchanan].—Saramma,

1963:77 [Kerala coast].

DESCRIPTION.—Based on 40 specimens, 90-138 mm

SL, including the holotypes and paratypes of C.

macrostomus and C. luctuosus.

Depth of body 23.11-28.11 (M = 25.89), length of

head 23.97-30.83 (M = 26.39) percent of standard

length. Eye diameter 5.56-10.34 (M = 7.59), interorbital space sometimes absent, 1.39-4.17 (M = 2.94)

percent of length of head. Two nostrils on ocular

side, anterior nostril tubular, in front of lower eye,

posterior nostril simple, in the anterior half of

interorbital space. Snout obtusely pointed, 20.59-

30.0 (M = 25.80) percent of length of head, rostral

hook short, extending to front of anterior nostril.

Maxillary extending to well beyond posterior margin of fixed eye; angle of mouth extending to

below vertical from posterior border of fixed eye

or just beyond it, nearer to tip of snout to branchial

opening, tip of snout to angle of mouth 35.29-47.50

(M = 40.49), angle of mouth to branchial opening

43.55-62.07 (M = 57.48) percent of length of head.

Scales: Ctenoid on both sides.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line curving onto dorsal fin at

a short distance from caudal base, midlateral line

with 80-92 (M = 83) scales; 14-16 (M = 15) scales

between them. No lateral line on blind side.

Interlinear scale rows

Frequencies

14 15 16

13 17 10

Fins: Dorsal with 100-106 (M = 103) rays, anal

with 78-84 (M = 80), caudal 10 in 23 specimens

(radiographs).

Vertebrae: 48-51 comprising 9 abdominal and

38-42 caudal elements in 23 specimens (radiographs).

Coloration: Upper side light brownish with dark

brown mottling on it, giving the appearance of

several irregular transverse bands, lower whitish in

preserved specimens.

Size: Largest specimen examined, 154 (138 + 18),

is from Cochin.

DISTRIBUTION.—Seas and estuaries of India; recorded from a depth of 8-12 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus macrostomus closely resembles C. semifasciatus, particularly with regard to the extension of the maxillary

beyond the posterior border of the fixed eye, the

relatively larger eyes, and the vertebral and fin-ray

counts. Cynoglossus macrostomus is, however, easily

distinguishable by the large number of interlinear

scale rows (14-16, M = 15 cf. 11-14, M = 12) and

the midlateral scale rows (80-92, M = 83 cf. 70-78,

M = 75), the more elongate body (depth 23.11-

28.33, M = 25.89 percent of SL cf. 24.53-31.58, M

= 27.88 percent), longer head (23.97-30.83, M =

26.89 percent of SL cf. 19.81-29.41, M = 22.76), and

the coloration, especially the distinctive blackish

nature of the fins.

NOTE ON SYNONYMY.—Norman (1928:204) considered C. hamiltonii described by Day (1877:436)

as distinct from C. hamiltonii Gunther (1862:504)

and described it as C. macrostomus, basing his

description on two specimens from "Calcutta and

Orissa." He characterized the new species as having

two lateral lines on ocular side, divided by 15 or

16 rows of scales. This species, which forms an

important commercial fishery on the west coast of

India and popularly known as the "Malabar sole,"

has all along been confused with C. semifasciatus

(Menon, 1971). Cynoglossus macrostomus can, however, be readily distinguished from C. semifasciatus

by its larger number of interlinear scales, 14-16
NUMBER 2S8 73

FIGURE 34.—Outline drawing of C. macrostomus (BMNH 1932.2.6.1) from Tanur, Malabar coast.

(M = 15) cf. 11-14 (M = 12). Another species confused with the Malabar sole is C. luctuosus Chabanaud (1947c:813) described on the basis of nine

specimens from Tanur on the Malabar coast and

characterized with 14-15 rows of scales between the

lateral lines on the ocular side. I have examined the

type of this species in London and found the species to be identical with C. macrostomus. I have,

therefore, synonymized them.

TYPE SPECIMENS.—Holotype of C. macrostomus,

ZSI 1460, 121 mm SL, Hooghly estuary near Calcutta, coll. F. Day; paratype, BMNH 1889.2.1.4074,

122.5 mm SL, Orissa, India, coll. F. Day. Holotype

of C. luctuosus, BMNH 1932.2.6.8, 133 mm SL,

Tanur, coll. D. W. Devanasen; 8 paratypes, BMNH

1932.2.6.1-7 and 9, 106-138 mm SL, Tanur obtained

along with holotype.

OTHER MATERIAL EXAMINED.—INDIA: 1 specimen, BMNH

1928.3.20.132, Portugese India, coll. S. W. Kemp. 2, ZSI 143,

Mormugao Bay, Goa, coll. S. W. Kemp. 4, ZSI 173-6, Mormugao Bay, Goa, coll. S. W. Kemp. 1, SOSC Ref 334,

Neendakara, Kerala, lat. 08°55'N, long. 76°30'E, coll. Anton

Bruun, 6.10.1966. 1, SOSC Ref 334, Ernakulam, Kerala,

lat. 10°0'N, long. 76°12'E, coll. Anton Bruun. 1, SOSC

Ref 334, Kerala, lat. 09°2rN, long 76°17'E, coll. Anton

Bruun. 22, SOSC Ref 334, Neendakara, Kerala, lat. 08°56'N,

long. 76'30'E, coll. Anton Bruun. 17, SOSC Ref 334,

Cochin, Kerala, lat. 10°00'N, long. 76°08'E, coll. Anton

Bruun. 1, ANSP 74855, Calicut, coll. Madras Fisheries

Department.

The monopus complex

Cynoglossus monopus, the only species of the

monopus complex, is typified by its small pedunculate eyes placed close together before the eyes. Its

range extends from the Malay Archipelago to seas

of India toward the west, and northward along the

South China Sea to Hong Kong.

32. Cynoglossus monopus (Bleeker)

FIGURE 35; PLATE 14

Plagusia monopus Bleeker, 1849:11 [type-locality: Bali].

Cynoglossus monopus.—Bleeker, 1875:38, pi. 245: fig. 4.—

Alcock, 1896:330.—Norman, 1928:204.—Weber and de

Beaufort, 1929:197.—Chu, 1931:94.—Smith, 1933:84.—

Fowler, 1934b:219.—Suvatti, 1936:94; 1950:326.—Kuronuma,

1961:32.—Punpoka, 1964:66 [Sumat Prakarn Province,

Thailand].

Plagusia melanopterus Bleeker, 1851a:415; 1852:25.

Arelia melanopterus.—Bleeker, 1859:184.

Cynoglossus melanopterus.—Gunther, 1862:502.—Alcock, 1889:

289—Seale, 1914:78 [Hong Kong].—Chu, 1931:94.

Arelia ceratophrys Kaup, 1858:108.

DESCRIPTION.—Based on 11 specimens, 94.0-153.0

mmSL.

Depth of body 21.90-26.12 (M = 23.58), length of

head 19.46-21.57 (M = 20.37) percent of standard

length. Eyes small, close together and pedunculate;

diameter of eye 4.0-6.78 (M = 5.80) percent of

length of head, interorbital space almost absent.

Two nostrils on ocular side, close together in front

of the anterior border of fixed eye, anterior nostril

tubular, posterior simple. Snout obtusely pointed,

26.32-37.29 (M = 31.82) percent of head, rostral

hook rather short, ending just in front of anterior

nostril. Maxillary extending to below posterior border of fixed eye or just behind it; angle of mouth

extending to below vertical from posterior half of

fixed eye, nearer to tip of snout than to branchial

opening; tip of snout to angle of mouth 39.39-47.62

(M = 44.06), angle of mouth to branchial opening

53.06-64.41 (M = 57.76) percent of length of head.

Scales: Ctenoid on both sides, including those

on lateral lines.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line undulated, entering dorsal
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 35.—Outline drawings of C. monopus (BMNH 1862.6J.13) from Bleeker's collection.

fin along 3rd to 5th ray, counted from the rear,

midlateral line with 108-125 (M = 117) scales, 16-

20 (M = 17) scales between them. No lateral line

on blind side.

Interlinear scale rows 16 17 18 19 20

Frequencies 5 2 1 2 1

Fins: Dorsal with 115-120 (M = 117) rays, anal

with 92-96 (M = 94), caudal 10 in 6 specimens

(radiographs).

Vertebrae: 55-56, comprising 9 abdominal and

44-47 caudal elements in 6 specimens (radiographs).

Coloration: Upper side light brownish, lower

whitish in preserved specimens.

Size: Largest specimen examined, 188.5 mm in

SL, is from Bangkok.

DISTRIBUTION: From Malay Archipelago to Hong

Kong and to northern part of Bay of Bengal;

recorded from a depth of 7-10 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus monopus

occupies a rather isolated position with regard to

the position of the nostrils, which are placed close

together in front of the anterior border of the fixed

eye, and the small pedunculate eyes. The most

closely related species is C. macrostomus. In the

obtusely pointed snout, the elongate nature of the

body, and especially in the blackish coloration of

the posterior portion of the fins in fresh specimens,

these species resemble each other to a certain extent.

There is no doubt that C. monopus evolved from a

macrostomus-Uke ancestor.

NOTE ON SYNONYMY.—Norman (1928:204) included P. melanopterus Bleeker and A. ceratophrys

Kaup in the synonymy of C. monopus (Bleeker) and

I concur with him.

TYPE SPECIMENS.—Bleeker (1849:11) described P.

monopus from Batavia and Boleling, Bali and

another species P. melanopterus Bleeker (1851a:415)

from Batavia and Boleling, Bali, actually as a

replacement name for P. monopus and obviously

based on the same types, but possibly including a

few additional specimens for P. melanopterus collected after the publication of P. monopus but
NUMBER 238 75

earlier than that of C. melanopterus. Unfortunately,

Bleeker did not mention the number of specimens

on which the description of P. monopus was based

but only the maximum size of the specimens, namely

180 mm, was indicated. This maximum size mentioned for P. monopus is obviously erroneous as

in the subsequent description of P. melanopterus

consisting of the same material. The maximum size

mentioned is 170 mm, Bleeker's (1850a:415) description of P. melanopterus being based on 11 specimens, 95 to 170 mm in TL.

According to Dr. M. Boeseman, the specimens

cataloged as RMNH 6799 (1879 auction) originally

contained 11 specimens, including the types of both

P. melanopterus and P. monopus, but there is no

specimen of 170 mm in length in it, and there

may or may not be a syntype of P. monopus in the

original material. According to him, however, it

seems reasonable to assume that actually the maximum size must have been only 160 mm, as the jar

originally included 11 examples, 83-147 (92.5-159)

mm.

Chabanaud in 1946 removed two of the smaller,

83-100 (92.5-100) mm, specimens to C. oxyrhynchus

and a specimen of 130 (141) mm was selected as

the lectotype of C. melanopterus and indicated as

such in the label.

Since C. melanopterus was described as a replacement name for C. monopus, it is appropriate to

consider the 141 mm specimen as the lectotype of

P. monopus. It is accordingly selected here, retaining

RMNH 6799. The remaining 8 specimens are

paralectotypes of both C. melanopterus and C.

monopus are recataloged as RMNH 26212.

OTHER SPECIMENS EXAMINED.—INDONESIA: 1 specimen,

BMNH 1862.6.3.13, loc. not given, coll. P. Bleeker. 2,

BMNH 1931.4.23.43-44, Java, coll. Hardenberg. 1, BMNH

1931.4.23-45, Bay of Batavia, coll. Hardenberg. 3, AMNH

17014, Sumatra, coll. Zeeviss, 1941. 6, AMNH 19815,

Sumatra, coll. Zeeviss, 1941. 1, ZMC. (unregistered), St.

Helena, coll. Th. Mortensen, 1930. PHILIPPINES: 1.

AMNH 19645, Celebes, coll. Zeeviss, 1941. THAILAND: 1,

USNM 88374, Thailand, coll. H. M. Smith. 9, CAS (GVF),

Samut Prakan Province, Gulf of Thailand, coll. Thai Fisheries. 1, ANSP 59919, Bangkok, coll. R. M. de Schaunensee.

SINGAPORE: 1, MCZ 33209, Singapore. CEYLON: 2.

BMNH 1933.12.27.1-2, Ceylon, coll. Harvard. INDIA: 3,

BMNH 1890.12.4.14-16, Ganjam coast, ex. Indian Museum.

2, BMNH 1928.3.20.99-100, Orissa coast, ex. Indian Museum. 2, SOSC, Porto Novo, Madras, lat. 11°35'N, long.

79°50'E, coll. Anton Bruun, 1966. 1, SOSC, Bombay,

Maharastra, lat. 18"50'N, long. 72°35'E, coll. Anton Bruun.

1, SOSC, Porto Novo, Madras, lat. ll°30'N, long. 79"WE.

coll. Anton Bruun. 6, ZSI 124.20-25, Orissa coast, 7 fms,

coll. Investigator. 1, ZSI 12617, Ganjam coast, coll.

Investigator.

The puncticeps complex

The puncticeps complex has its center of distribution in the Indo-Australian Archipelago including northwest Australia, Philippines, the seas of

India and Pakistan, and farther westward to the

East Coast of Africa. This complex is closely related

to the cynoglossus complex and both may have descended from a common ancestral stock.

The principal character that distinguishes this

complex is the smaller size of its body scales, the

interlinear scale rows being 14 to 19.

Included in this complex are C. puncticeps, C.

durbanensis, and C. gilchristi.

33. Cynoglossus puncticeps (Richardson)

FIGURE 36; PLATE 15

Plagusia puncticeps Richardson, 1846:280 [type-locality:

China].—Whitehead, 1969:218, pi. 29c.

Cynoglossus puncticeps.—Gunther, 1862:500.—Bleeker, 1875:

37. pi. 245: 15, fig. 7.—Day, 1877:437, pi. 97: fig. 1;

1889:459.—Alcock, 1889:289.—Jordan and Seale, 1907:46.—

Jenkins, 1910:30.—Norman, 1928:205.—Weber and de

Beaufort, 1929:198.—Chu, 1931:94.—Herre. 1932:433.—Wu,

1932:151 [Amoy, Pehai, Canton, Hong Kong, and

Hainan].—Smith, 1933:84.—Fowler, 1934b:220, fig. 34.—

Suvatti, 1950:328.—Herre, 1953:190.—Munro. 1955:265, pi.

51: fig. 771.—Fowler, 1956:137.—De Silva, 1956:198

[Panadura].—Munro, 1958:285 [Kau Kau].—Kuronuma,

1961:32—Punpoka, 1964:69 [Thailand].—Pradhan, 1964:

458.—Chen and Weng, 1965:91, fig. 62 [Tainan and

Tungkong].—Shen, 1967:214, figs. 138-141 [Lamma

Island].

Plagusia nigrolabeculata Richardson, 1846:280 [coasts of

China. Canton].—Kaup. 1858:110.—Whitehead. 1969:218,

pi. 29b.

Cynoglossus nigrolabeculatus.—Bleeker, 1873:131 [reference].

Plagusia aurolimbata Richardson, 1846:280 [coasts of

China].—Kaup. 1858:110.—Whitehead, 1969:218, pi. 286.

Cynoglossus aurolineatus.—Bleeker, 1873:130.

Plagusia javanica Bleeker, 1851a:414 [type-locality: Batavia];

1852:24.

Arelia javanica.—Bleeker, 1859:184.

Plagusia brachyrhynchos Bleeker, 1851a:414 [type-locality:

Batavia]; 1852:24.

Arelia brachyrhynchos.—Bleeker, 1859:184.

Cynoglossus brachyrhynchus.—Gunther, 1862:499.—Bleeker,

1875:37, pi. 243: fig. 4.—Day, 1877:435, pi. 96: fig. 4;

1839:457.—Johnstone, 1904:206.—Weber, 1913b:443.

Cynoglossus brevis Gunther, 1862:500 [type-locality:
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ganges].—Day, 1877:437, pi. 97: fig. 2.—Alcock, 1889:289.—

Hora, 1923b:760.—Norman, 1928:206.

Cynoglossus lida [not Bleeker].—Jenkins, 1910a:31.

Cynoglossus lida var. punctatus.—Jenkins, 1910a:30.

Cynoglossus puncticeps immaculata Pellegrin and Chevey,

1940:154 [Vietnam].

DESCRIPTION: Based on 87 specimens, 35.0-166.0

mm SL, including the paralectotype of C. brevis.

Depth of body 12.05-45.16 (M = 28.43), length

of head 11.45-30.65 (M = 22.22) percent of standard length. Diameter of eye 6.25-16.13 (M = 10.71),

interorbital space rather narrow, 0.36-6.67 (M =

3.03) percent of length of head. Two nostrils on

ocular side, anterior nostrils tubular, in front of

lower eye, posterior nostril simple, immediately in

front of interorbital space. Snout rounded or obtusely pointed, 24.39-41.18 (M = 32.67) percent of

length of head, rostral hook short, usually ends in

front of anterior nostril, rarely reaching below

anterior border of fixed eye. Maxillary extending to

FIGURE 36.—Lateral view of ocular side of three specimens of C. puncticeps showing variation

in pigment pattern: a, specimen 143.5 mm SL (ZSI.F 5358/2) from Alleppey, Kerala; b#,

specimens 110.6 and 1353 mm SL from Cochin coll. R. R. C. Edwards, Aberdeen.
NUMBER 238 77

below middle or posterior half of fixed eye; angle

of mouth extending to below vertical from anterior

half or middle of fixed eye, usually nearer to tip

of snout than to branchial opening; tip of snout

to angle of mouth 34.21-51.43 (M = 44.16), angle

of mouth to branchial opening 41.67-72.73 (M =

55.62) percent of length of head.

Scales: Ctenoid on both sides, including those of

lateral lines.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line slightly undulating, running

backward and entering dorsal fin along 5th-6th ray,

counted from the rear, midlateral line with 78-99

(M = 87) scales, 14-19 (M = 16) scales between

them. No lateral line on blind side.

14 15 16 17 18 19

4 6 22 23 16 9

Interlinear scale rows

Frequencies

Fins: Dorsal with 90-100 (M = 96) rays, anal with

72-78 (M = 75) rays, caudal 10 in 57 specimens

(radiographs).

Vertebrae: 44-49, comprising 9 abdominal and

35-40 caudal elements in 57 specimens (radiographs).

Coloration: Upper side brownish with dark

blotches all over the body, appearing as somewhat

irregular crossbands that disappear with age, lower

whitish. Some of the rays of the vertical fins marked

with dark brown.

Size: Largest specimen examined, 175 (161 + 14)

mm, is from Sind, Pakistan.

DISTRIBUTION.—From northwest Australia, Malay

Archipelago to Philippines, through South China

Sea to Taiwan, and westward to seas of India;

recorded from a depth of 7-75 fms and known from

brackish waters as well.

DIAGNOSIS AND AFFINITIES.—The color pattern

with large irregular blotches on the head and body,

which are often united to form irregular crossbands,

and some of the rays of the vertical fins with dark

brown predominating are characteristic. Considerable variation from this pattern occurs, however,

probably with changes in the nature of the substratum or intensity of light penetration. Cynoglossus puncticeps, which is replaced by C.

durbanensis and C. gilchristi along the East Coast

of Africa, is distinguished by 10 caudal fin rays

whereas in the other species the number is only

8.

NOTE ON SYNONYMY.—Norman (1928:205) included P. javanica Bleeker and P. brachyrhynchos

Bleeker in the synonymy of C. puncticeps and I

concur with him. The additional names included

here are P. nigrolabiculata Richardson, P. aurolimbata Richardson, C. brevis Giinther, and C.

p. immaculata Pellegrin and Chevey.

Richardson's descriptions of P. puncticeps, P.

aurolimbata, and P. nigrolabiculata were all based

on Reeves Chinese fish drawings (see Whitehead,

1969). Fowler (1934b) synonymized the latter two

species with C. puncticeps. I have examined Reeve's

original drawings of these species in the British

Museum and found no significant difference

between them.

Gunther (1862:500) described C. brevis on the

basis of a single specimen, 108 mm in TL, from the

Ganges and characterized it with two lateral lines

on left side separated by 17 scale rows. I have

examined the holotype of C. brevis in the British

Museum and I am convinced that differences in

the proportional measurements such as the longer

head length and longer snout in the type are

attributable to intraspecific variations and that the

two species are essentially the same in all respects

including the color pattern, though considerable

variation in pigmentation with age and change in

the substratum is exhibited by the fish.

Pellegrin and Chevey described C. p. immaculata

on the basis of a specimen from Bac Lieu, Cochin

China. I have examined the holotype of C. p.

immaculata in Paris and found it to exhibit no

significant difference with C. puncticeps; C. p.

immaculata is, therefore, synonymized here with

C. puncticeps.

TYPE SPECIMENS.—Bleeker's description of P.

javanica was based on 17 specimens, 100-155 mm

in TL. In the Bleeker collection in Leiden, cataloged

as RMNH 6797, there are 20 specimens as C.

puncticeps, obviously the specimens of P. javanica

having been added to C. puncticeps after its

synonymy. Out of the 20 specimens, 17 specimens,

90-109 (100-120) mm, are types and 3 specimens,

83-84 (91-92) mm, are nontypes. None of these

agree with Bleeker's upper size limit of 155 mm,

which may well be erroneous. Dr. Boeseman examined the material on my behalf and selected a

specimen of 90 mm SL, 100 mm TL, as the lectotype of P. javanica. It was retained as RMNH 6797

and the remaining 16 specimens were removed as

paralectotypes of P. javanicus and recataloged as

RMNH 26206.
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Bleeker's (1851a:414) description of P. brachyrhynchos was based on two specimens, 118 and 122

mm in TL, from Batavia. In the Bleeker collection

specimens in Leiden cataloged as RMNH 6796,

there are nine specimens one of which, 114 mm

SL, 123 mm in TL, from Batavia, is selected as

the lectotype of P. brachyrhynchos. The lectotype

is retained as RMNH 6796 and the paralectotypes

of 52-145 (56-? caudal mutilated) mm have been

recataloged as RMNH 26207.

Holotype of C. brevis, BMNH 1855.12.26.602,

80 mm SL, from Ganges, ex. Zool. Society.

Holotype of Cynoglossus puncticeps immaculate

Pellegrin and Chevey, MNHP 40.39, from Bac Lieu,

Cochin China.

OTHER MATERIAL EXAMINED.—CHINA: 2 specimens, BMNH

1848.3.16 I94a,b, China. 2, BMNH 1850.3.19.916 a,b, China.

I, BMNH 1851.12.27.215. China. 1, BMNH 1851.12.27.344.

China. 1, BMNH 1924.12.15.67, Hainan Is., coll. Light. 4,

BMNH 1924.12.15.68-71, Amoy. coll. Light. 10. BMNH

1924.12.15.72-81, Santuao, coll. Light. 9. BMNH 1924.12.

15.80 a-h, Santuao. coll. Light. 1, BMNH 1930.7.28.16.

Zingkow. coll. Chen. 2. BMNH 1930.7.28.14-15, Haikon,

coll. Chen. 1, BMNH 1936.10.7.57, Foochow, coll. Chen. 4,

USNM 6937, 6958. China. 1. UMMZ 70359. China, coll.

Bead and Sleeve.

PHILIPPINES: 1, BMNH 1872.10.18.133. Manila, coll.

Meyer. 1. BMNH 1879.5.14.82. Philippines, coll. Challenger.

1, USNM 83075. Philippines. 1. USNM 1068026, Philippines,

coll. H. C. Keller. 16, USNM 112871, 78-81. Philippines,

coll. H. C. Keller. 1, USNM 113179, Philippines, coll.

H. C. Keller. 18, USNM 137695-137703, Philippines, coll.

H. C. Keller. 18. USNM 137695-137703, Philippines, coll.

Albatross. 1, ANSP 635524, Philippines, coll. J. Clemense. 8,

ANSP 82548, Philippines, coll. J. Clemense. 1. ANSP

82547, Philippines, coll. J. Clemense.

SINGAPORE: 1. BMNH 1933.12.27.6, Singapore, ex.

Indian Museum. 2, BMNH 1933.7.31-32. Singapore, ex.

Raffles Museum. 2, BMNH 1934.9.6.5-6, Singapore Mkt.,

ex. Raffles Museum. 10, BMNH 1934.1121.9-18, Singlap, ex.

Raffles Museum. 3, MCZ 11510. Singapore.

THAILAND: 1, USNM 109799, Gulf of Thailand, coll.

H. M. Smith. 4, ANSP 61837, Paknam. 5, ANSP 61843-55,

Paknam, coll. R. de Schanenam. 1, ANSP 61842, Paknam,

coll. R. de Schanenam. 2, ANSP 61852-53, Bangkok, coll.

R. de Schanenam. 1. ANSP 61841, Bangkok, coll. R. de

Schanenam. 2. ANSP 61852-53. Bangkok. coD. R. de

Schanenam. 1, ANSP 61841, Bangkok, coll. R. de Schanenam.

1. ANSP 61854, Bangkok, coll. R. de Schanenam. 2. ANSP

87437, Bangkok, coll. R. de Schanenam. 1, CAS (GVF),

Gulf of Thailand, coll. Naga Expd.

BORNEO: 1, BMNH 1894.1.19.1932, Bruntal (Sarawak),

coll. Brooke.

INDONESIA: 1, BMNH 1931.4.21.47, Java, coll.

Hardenberg. 1. BMNH 1931.4.23.46, Batavia Mkt., coll.

Hardenberg. 1. ZMA 108.002, Indonesia, coll. Tydeman. 5,

UMMZ (unregistered), Java. 2, AMNH 18414, Java. 1.

NHV 43806, Java, coll. E. Novara. BURMA: 1. ZSI 1462,

Moulmein, coll. F. Day.

INDIA: 1, BMNH 1852.9.13.235 B, Ganges, ex. Zool.

Soc. 1, BMNH 1855.9.19.47, locality unknown, coll. Haslar.

1, BMNH 1862.6.3.15, locality unknown, coll. P. Bleeker. 1,

BMNH 1862.6.3.20, locality unknown, coll. P. Bleeker. 1,

BMNH 1889.2.1.4069, Bombay, coll. F. Day. 1. BMNH

18892.1.4072, Andamans, coll. F. Day. 1, BMNH 1889.2.1.

4075, Calcutta, coll. F. Day. 1, BMNH 1928.3.20.113, Trivandrum, ex. Indian Museum. 1, BMNH 1928.3.20.114.

Trivandrum, Travancore, ex. Indian Museum. 4, BMNH

1928.3.20.115-118, Ganjam coast, ex. Indian Museum. 2,

BMNH 1928.320.119-120, Ganjam coast, 7-14 fms, ex.

Indian Museum. 2, BMNH 1928.320.121-122. Chilka Lake,

ex. Indian Museum. 1, BMNH 1928.3.20.123. Chilka Lake,

ex. Indian Museum. 5, BMNH 1928.3.20.124-128, Chilka

Lake, ex. Indian Museum. 2, BMNH 1928.320.129-30, Orissa

coast, 7 fms, ex. Indian Museum. 1, BMNH 1928.320.131,

Ganjam coast, 8-9 fms, ex. Indian Museum. 1, BMNH

1933.12-7, Calicut, Malabar, coll. Devanesan. 2, BMNH

1938.8.9.11.12. Calicut, coll. Devanesan. 1. ZSI 139.

Travancore, coll. F. Day. 2, ZSI 161 and 282, Goa, coll.

5. W. Kemp. 1, ZSI 463, Calcutta bazar, ex. Asiatic Society.

1. ZSI 1147. Calicut, coll. F. Day. I, ZSI 1148, Bombay, coll.

F. Day. 1, ZSI 1459, Calcutta, coll. F. Day. 1, ZSI 2691,

Calcutta, coll. F. Day. 1, ZSI 2693, Madras, coll. F. Day. 2.

ZSI 12167, Orissa coast, coll. Marine Survey. 2, ZSI 12235

and 12673, Ganjam coast, Orissa, coll. Marine Survey. 8,

ZSI 12658-63 to 70, Ganjam coast, coll. Marine Survey. 1,

ZSI 12674, Ganjam coast, 4-14 fms, coll. Marine Survey. 2.

F 4188

ZSI - , Trivandrum, ex. Trivandrum Museum. 2. ZSI

F 857 9

 , Calcutta bazar, ex. Asiatic Society. 1, ZSI

 F 858 0

 , CalF 099

cutta bazar, ex. Asiatic Society. 1, ZSI , 5 mi SE of

Geokale, Patashmipur, Chilka Lake, Orissa. coll. K. S. Misra.

F 962

6. ZSI——=-, S of Cherryakuda Is., 4 mi NE of Ramba,

Chilka Lake, Orissa, coll. K. S. Misra. 5, ZSI L 165 5 , North

2

Ernakulam Back Waters, coll. F. H. Gravely. 10, ZSI

(unregistered), Kerala, coll. A. G. K. Menon. 9, ZSI

(unregistered), Barkul, Chilka Lake, coll. Chilka Survey,

1.3.1914. 1, ZSI (unregistered). Barkul, Chilka, coll. Chilka

Survey, 5.3.1914. 2. ZSI (unregistered), S of Kalidai. Chilka

Lake, coll. Chilka Survey, 5.3.1941. 1, ZSI (unregistered),

Kalidai, Chilka Lake, coll. Chilka Survey, 21.2.1941. 14, ZSI

(unregistered), Channel off Barhampore, Chilka Lake, coll.

Chilka Survey, 2.9.1914. 1. ZSI (unregistered), Gopkuda

Bay, Chilka Lake, coll. Chilka Survey, 14.2.1914. 5, ZSI

(unregistered), Channel of Satpara, coll. Chilka Survey. 1,

ZSI (unregistered), Kalupara Ghat, Chilka Lake, coll. Chilka

Survey. 1, ZSI (unregistered), Patashnipur, Chilka Lake, coll.

Chilka Survey, 12.3.1914. 1, ZSI (unregistered), Long Is.,

near Manipatna, Chilka Lake, coll. Chilka Survey. 1, ZSI

(unregistered), Cheriya Is. to Barkuda Is. across mouth of

Rambha Bay, coll. Chilka Survey, 1914. 1, ZSI (unregistered),

Barkuda Is., Chilka Lake, coll. Chilka Survey, 172.1914. 1,

ZSI (unregistered), Cheriya Is. Bay, Chilka Lake, coll. Chilka
NUMBER 238 79

Survey. 1. ZSI (unregistered), channel between Satpara and

Barikuda, Chilka Lake, coll. Chilka Survey. 4, ZSI (unregistered), W of Satpara, Chilka Lake, coll. Chilka Survey. 2.

ZSI (unregistered), Cochin, coll. Edward. 4, ZSI (unregistered), Cochin, coll. Dr. A. Daniel, 22.9.1960. 2. ZSI (unregistered), Cochin, coll. Dr. Quasim. 1, ZSI (unregistered),

Aleppy, Kerala, coll. Chakrapany. 1, ZSI (unregistered),

Travancore, coll. Dr. £. G. Silas. 1, ZSI (unregistered),

Karwar, coll. K. K. Tiwari. 1, SOSC, Madras, coll. Anton

Bruun. 1, ANSP 74855, Calcutta. 1. NHV 43770. Calcutta,

coll. F. Day.

PAKISTAN: 1. BMNH 1889.2.1.4068, Sind. coll. F. Day. 2,

BMNH 1911.12.6.17.18, Karachi, coll. Townsend. 1, NHV

43772. Sind. coll. F. Day.

34. Cynoglossus durbanensis Regan

FICURE 37; PLATE 15

Cynoglossus durbanensis Regan, 1921a:2 [type-locality:

Durban. South Africa].—Bonde, 1922:25.—Barnard, 1925:

412.—Smith. 1949:165.

DESCRIPTION.—Based on 7 specimens, 88.5-182.0

mm SL, including the lectotype and paralectotype

of C. durbanensis.

Depth of body, 25.82-30.88 (M = 28.31), length

of head 17.43-20.90 (M = 18.95) percent of standard

length. Diameter of eye 8.20-10.87 (M = 9.78), interorbital space 6.38-10.81 (M = 8.67) percent of

length of head. Two nostrils on ocular side, anterior

nostril tubular, in front of lower eye, posterior nostril simple, in front of interorbital space. Snout obtusely rounded, 23.40-33.96 (M = 30.20) percent of

length of head, rostral hook short, ends in front of

anterior nostril. Maxillary extending to below posterior half of fixed eye; angle of mouth extending

below vertical from middle or posterior half of fixed

eye, nearer to tip of snout than to branchial opening; tip of snout to angle of mouth 40.43-49.06

(M = 43.70), angle of mouth to branchial opening

55.32-63.04 (M = 58.11) percent of length of head.

Scales: Ctenoid on both sides, including those of

lateral lines.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line usually entering dorsal fin

along 3rd-5th ray counted from the rear, midlateral

line with 90-106 (M = 99) scales, 18-21 (M = 20)

scales between them. No lateral line on blind side.

Interlinear scale rows

Frequencies

18 19 20 21

1 1 4 1

Fins: Dorsal with 98-105 (M • 102) rays, anal

with 78-84 (M = 81) rays, caudal 8(9) in 5 specimens (radiographs).

Vertebrae: 48-50, comprising 9 abdominal and

39-41 caudal elements in 5 specimens (radiographs).

Coloration: Light brownish with darker irregular

blotches and dots in preserved material.

Size; Largest specimen examined (lectotype) is

184 (170 + 14) mm from Durban.

DISTRIBUTION.—Africa (Natal to Zanzibar).

DIAGNOSIS AND AFFINITIES.—The nearest relative

of C. durbanensis is C. gilchristi. The features that

distinguish this species are: the deeper body (mean

depth 28.31 percent of SL cf. 27.16 percent), smaller

eyes (mean diameter 9.78 percent of head cf. 11.56

percent), wider interorbital space (mean width 8.67

percent of head cf. 6.22 percent), and smaller body

scales, the interlinear scale rows being 18-21, M =

19, whereas in C. gilchristi there are only 14-15,

M = 14.

TYPE SPECIMENS.—BMNH 1920.7.23.37, 170 mm

FIGURE 37.—Outline drawing of C. durbanensis (BMNH 19223.5.20) from Lumbo.
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SL, from Durban, coll. H. W. B. Marley, an excellent example of the species, is selected here as the

lectotype of C. durbanensis and BMNH 1920.7.23.

38, 152 mm SL, collected along with the lectotype

as the paralectotype.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH 1920.12.

6.S0, Durban, ex. Transvaal Museum. 1, BMNH 1922.5.550,

Lumbo, E Africa, coll. Laveridge. 1, ANSP 63890, Durban,

coll. H. W. Bell Marley, 1935. 1, ANSP 87113, Natal coast,

coll. H. W. Bell Marley, 1935. 3, MCZ 11519, Zanzibar, coll.

Cooke.

35. Cynoglossus gilchristi Regan

FIGURE 38; PLATE 16

Cynoglossus brachycephalus [not Bleeker].—Gilchrist, 1905:12,

pi. 30.—Bonde, 1922:25.—Gilchrist and Thompson,

1917:399.

Cynoglossus gilchristi Regan, 1920:222 [type-locality: Natal,

off mouth of Umhlanga River].—Barnard, 1925:412.—

Bonde, 1925:294.

Cynoglossoides gilchristi.—Smith, 1949:165, fig. 337.

DESCRIPTION.—Based on 4 specimens, 44.0-133.0

mm SL, including the holotype of C. gilchristi.

Depth of body 24.81-28.73 (M = 27.16), length of

head 18.0-23.86 (M = 21.58) percent of standard

length. Diameter of eye 8.0-18.18 (M = 11.56),

interorbital space 4.55-7.89 (M = 6.22) percent of

length of head. Two nostrils on ocular side, anterior

nostril tubular, in front of lower eye, posterior nostril simple, in front of the interorbital space. Snout

rounded, 26.0-31.82 (M = 28.84) percent of length

of head, rostral hook rather short, ends below

anterior nostril or just in front of it. Maxillary

extending to below middle or posterior half of fixed

eye; angle of mouth extending to below vertical

from anterior half or middle of fixed eye, nearer

to tip of snout than to branchial opening; tip of

snout to angle of mouth 44.0-47.62 (M = 46.11),

angle of mouth to branchial opening 54.55-57.89

(M = 56.40) percent of length of head.

Scales: Ctenoid on both sides, including those of

lateral lines.

Lateral-Line System: Two lateral lines on ocular

side, dorsolateral line more or less undulated,

generally stopping at four-fifths of length of the

body and becoming obscure; midlateral line with

74-80 (M = 77) scales, 14-15 (M = 15) scales between them. No lateral line on blind side.

Interlinear scale rows 14 15

Frequencies 1 S

Fins: Dorsal with 104-110 (M = 106) rays, anal

with 82-86 (M = 84) rays, caudal 8 in 3 specimens

(radiographs).

Vertebrae: 50 comprising of 9 abdominal, 49

caudal elements in 3 specimens (radiographs).

Coloration: Upper side light brown with irregular

darker blotches, lower whitish in preserved specimens.

Size: Largest specimen examined (holotype), 141

(133 + 8) mm, is from Natal.

DISTRIBUTION.—From Natal and Delagoa Bay to

Nossi Be, Madagascar; recorded from depths of

4-30 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus gilchristi

is closely related to C. durbanensis; the differences

distinguishing it from C. durbanensis are outlined

under C. durbanensis.

TYPE SPECIMENS.—Holotype of C. gilchristi,

BMNH 1903.9.29.2, 133 mm SL, Umlanga R., Natal,

S Africa, coll. Gilchrist.

FIGURE 38.—Outline drawing of C. gilchristi, holotype (BMNH 1903.9.29.2) from Natal.
NUMBER 238 81

OTHER MATERIAL EXAMINED.—3 specimens, SOSC Ref 134,

Nossi Be. Madagascar, lat. 1S°2O'5O''S, long. 48819'50"E,

coll. Te-vega Expd.

The lida complex

Cynoglossus lida, the only species forming the

lida complex, is characterized by a broadly round

prominent snout, the angle of the mouth being

distinctly nearer to the branchial opening than to

the tip of the snout and the rostral hook extending

to as far below the middle of the fixed eye. In all

other features it appears to be closely related to the

puncticeps complex and may have evolved from a

common ancestral stock.

36. Cynoglossus lida (Sleeker)

FIGURE 39; PLATE 16

Plagusia lida Bleeker, 1851a:413 [type-locality: Batavia];

1852:23.

Arelia lida.—Bleeker, 1859:184.

Cynoglossus lida.—Gunther, 1862:498.—Bleeker, 1875:36, pi.

243: fig. 2.—Day, 1877:436, pi. 97: fig. 3; 1889:458.—

Alcock. 1889:288.—Vindguerra, 1889:188.—Jenkins, 1910a:

30.—Regan, 1920:221, fig. 5 [Durban].—Bonde, 1922:25.—

Barnard, 1925:411 [Natal coast].—Fowler, 1925:187

[Natal].—Norman, 1928:210.—Weber and de Beaufort,

1929:203.—Smith, 1933:83.—Fowler, 1935:405 [Natal and

Zululand].—Smith, 1949:166.—Herre. 1953:190.—Munro,

1955:206.—Silva, 1956:199.—Fowler, 1956:187. fig. 102 —

Punpoka, 1964:61.—Chen and Weng, 1965:94 [Taiwan].

Plagusia polytaenia Bleeker, 1853c:529 [type-locality: Priaman].

Arelia polytaenia.—Meeker, 1859:185.

Cynoglossus polytaenia.—Bleeker, 1875:36, pi. 244: fig. 1.—

Weber and de Beaufort. 1929:201.

Cynoglossus intermedius Alcock, 1889:288.

Cynoglossus os Fowler, 1904:556 [type-locality: Padang,

Sumatra].

DESCRIPTION.—Based on 23 specimens, 45.0-200.0

mm SL, including the holotype of C. intermedius.

Depth of body 21.92-27.55 (M = 23.78), length of

head 21.15-25.56 (M = 23.30) percent of standard

length. Diameter of eye 3.70-8.89 (M = 6.32), interorbital space 3.19-7.41 (M = 3.93) percent of length

of head. Two nostrils on ocular side, anterior

nostril tubular, in front of lower eye, posterior

nostril simple, in anterior half of interorbital space.

Snout rounded, 30.43-43.33 (M = 37.99) percent of

length of head, rostral hook rather long, extending

to as far as below middle of fixed eye. Maxillary

extending to below posterior border of fixed eye,

or just behind it; angle of mouth extending to

below vertical from posterior half of fixed eye,

nearer to branchial opening than to tip of snout;

tip of snout to angle of mouth 44.44-57.45 (M =

53.03), angle of mouth to branchial opening 41.57-

55.56 (M = 46.55) percent of length of head.

Scales: Ctenoid on both sides.

Lateral-Line System: Two lateral lines on ocular

side, midlateral line with 72-90 (M = 80) scales,

12-15 (M = 13) scales between middle and upper

lateral line. No lateral line on blind side.

Interlinear scale rows 12 13 14 15

Frequencies \ 16 37 20 2

Fins: Dorsal with 99-108 (M = 105) rays, anal

with 77-85 (M = 81) rays, caudal 10 in 57 specimens (radiographs).

Vertebrae: 47-52, comprising 9 abdominal and

38-43 caudal elements in 57 specimens (radiographs).

Coloration: Upper side light brownish, lower

whitish in preserved specimens.

Size: Largest specimen examined is 213.0 (200 +

13) mm, trawled by Anton Bruun.

DISTRIBUTION.—From Malay-Archipelago and

I cm

FIGURE 39.—Outline drawing of C. lida (BMNH 1919.9.12.50) from Durban.
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Philippines through seas of India and Pakistan to

East Coast of Africa; recorded from a depth of

13-15 fms.

DIAGNOSIS AND AFFINITIES.—Superficially C. lida

resembles C. puncticeps in many morphometric

characters. It is distinguishable, however, by the

lack of dark irregular blotches on the body and

the dark brown marking of some of the rays of the

vertical fins and by the shape of its broadly rounded

snout (mean length 37.99 cf. 32.67 percent of head).

It is also characterized by the rictus of the mouth

being much nearer to the gill opening than to the

end of snout (mean distance from rictus of mouth

to gill opening 46.55 percent of head cf. 55.62

percent of head).

NOTE ON SYNONYMY.—Bleeker's description of P.

lida was on the basis of five specimens, 100-145 mm

in TL, from Batavia. He characterized it as having

two lateral lines on the left side separated by 13

series of scales and having the angle of the mouth

nearer to the gill opening than to the end of the

snout. Norman (1928:210) synonymized C. intermedius Alcock with C. lida. I have examined the

holotype of C. intermedius in the Zoological Survey

of India and find that the differences noted by

Alcock in the type of C. intermedius, such as the

more numerous, weakly ctenoid scales and the

longer head, are attributable to intraspecific variation and that they are conspecific. Gunther (1862:

498) considered P. polytaenia Bleeker (1853e:529)

conspecific with P. lida though Weber and de

Beaufort (1929:201) retained it as a separate species

from C. lida. I examined a specimen, BMNH 1862.

6.3.14, of P. polytaenia from the Bleeker collection

in London and found it to conform well with

Bleeker's description and figure of P. lida. Fowler

(1904:556) described C. os based on a single specimen from Pedang. I have not been able to locate

the type in Philadelphia (pers. comm., Dr. James

C. Tyler), but from the description it is found

to exhibit no significant difference from P. lida.

TYPE SPECIMENS.—Among Bleeker's material in

Leiden is one obviously composite lot (RMNH

6791) of 6 specimens, including the 5 syntypes of

P. lida (pers. comm., Dr. M. Boeseman). Since

Bleeker in his Atlas (1875:6, pi. 243: fig. 2) gave

the illustration of a specimen measuring 145 mm in

TL, obviously the largest specimens in the syntype

series, a specimen 134 mm SL, approximately 145

mm in TL, in the lot RMNH 6791, is selected as

lectotype of P. lida and retained in RMNH 6791.

The other five examples, 91-126 (100-136) mm,

recataloged as RMNH 26208, include 4 paralectotypes of P. lida.

Bleeker described P. polytaenia on the basis of

a single specimen, 107 mm in TL, from Priaman,

Sumatra. Among the Bleeker collection in Leiden

is a specimen 100 mm in SL, 107 mm in TL,

indicated as holotype, RMNH 6790, presumably by

Chabanaud (pers. comm., Dr. Boeseman).

Holotype of C. intermedius, ZSI 12246, 192.5 mm

SL, Orissa coast, coll. Marine Survey.

OTHER MATERIAL EXAMINED.—PHILIPPINES: 1 specimen,

BMNH 1872.4.6.96, Celebes, coll. Meyer. 12, USNM (unregistered), Philippines, coll. Albatross, 1907-09. 5, USNM

137955-59, Philippines, coll. Albatross. 9, USNM (unregistered), Philippines, coll. Albatross. INDONESIA: 1, BMNH

1862.6.3.14, locality not given, coll. P. Bleeker. 1, BMNH

1862.6.3.16, locality not given, coll. P. Bleeker. 1, ZMA OR276101, Indonesia. BURMA: 1, BMNH 1889.2.1.4064, Sittoung, coll. F. Day. 1, ZSI 2688, Burma, coll. F. Day.

INDIA: 1, BMNH 1928.3.20.129-30, Orissa coast, ex. Indian

Museum. 1, BMNH 1928.3.20.31, Ganjam coast, ex. Indian

Museum. 2, ZSI 12259-60, Orissa coast, coll. Marine Survey. 2, ZSI 12684-85, Orissa coast, coll. Marine Survey. 2,

ZSI F H p and F -^ , Tranquebar, E coast. India, coll.

194/5

A. G. K. Menon. 1. ZSI F -=~, Kilingimedu, Karaikkal,

IOOR

coll. A. G. K. Menon. 1, ZSI F •—, Sonakuppam, Cuddalore. coll. A. G. K. Menon. 28, SOSC Ref 170. Madras, coll.

Anton Bruun. AFRICA: 1, BMNH 1919.9.1230. Durban,

coll. Marley. 13, SOSC Ref 170. Mozambique coast, lat.

19°09'S, long. 36°3O'E, coll. Anton Bruun. 8, SOSC Ref 170,

NW coast of Madagascar, coll. Anton Bruun. 1, SOSC Ref

170, Mozambique coast, coll. Anton Bruun. 2, ANSP 64104-

05, Mozambique, coll. Albatross. 1, ANSP 84664, Natal, coll.

H. W. Bell Marley. 1, ANSP 86319, Natal, coll. H. W.

Bell Marley. 2, MCA 34108, Mozambique, coll. T. Barbous.

1, RU 275565, Inhaca.. coll. J. L. B. Smith.

The carpenteri group

This group is basically a natural assemblage of

specialized, lately evolved species of cynoglossus.

The principal characters distinguishing this group

are three lateral lines on the ocular side, none on

the blind side, two nostrils on ocular side, snout

long and pointed, with the angle of mouth situated

nearer to branchial opening than to tip of snout,

small eyes usually with a narrow interorbital space

(absent in C. suyeni), small scales, the interlinear

scale count being 15-22. The species of this group

are distributed on one hand in the Philippines
NUMBER 238 83

through Celebes to the Timor Sea and on the other

in the seas of India and Pakistan to the East Coast

of Africa. This group is most probably derived from

a cynoglossus-liine ancestor through elongation of

the body (especially the snout), reduction in the

size of the body scales, and addition of a ventrolateral line on the ocular side.

The carpenteri complex

Cynoglossus carpenteri, C. acutirostris, C. marleyi,

and C. suyeni form the carpenteri complex, which

is the only complex under the carpenteri group.

The characteristics of the complex are, therefore,

the same as given above for the group.

37. Cynoglossus carpenteri Alcock

FIGURE 40; PLATE 17

Cynoglossus carpenteri Alcock, 1889:287, pi. 18: fig. 1 [typelocality: Orissa coast]; 1890a:217; 1896:330; 1898. pi. 22:

fig. 5; 1899:133.—Boulenger, 1901:263.—Regan, 1905b:329.—

Norman, 1928:196.—Saramma, 1963:75 [Kerala coast].

DESCRIPTION.—Based on 21 specimens, 60.0-181.0

mm SL, including lectotype and paralectotype of

C. carpenteri.

Depth of body 23.81-29.5 (M = 26.66), length of

head 27.61-33.87 (M = 29.85) percent of standard

length. Diameter of eye 6.33-10.53 (M = 8.56),

interorbital space 2.08-6.56 (M = 3.74) percent of

length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior

nostril simple, in front of interorbital space. Snout

obtusely pointed, 30.85-36.84 (M = 33.32) percent

of head; rostral hook rather short, extends up to a

point from anterior nostril to anterior border of

fixed eye. Maxillary extending to well beyond fixed

eye; angle of mouth extending to below vertical

from posterior border of fixed eye or little beyond

it, either slightly nearer to branchial opening than

to tip of snout or midway between these two; tip

of snout to angle of mouth 39.02-56.10 (M =

50.76), angle of mouth to branchial opening 47.37-

54.55 (M = 48.78) percent of length of head.

Scales: Cycloid on ocular side with ctenoid scales

posteriorly, cycloid on blind side.

Lateral-Line System: Three lateral lines on ocular side, both dorsolateral and ventrolateral lines

usually entering dorsal and anal fins respectively

at a very short distance from caudal base, midlateral line with 75-96 scales, 15-19 (M = 16)

scales between them. No lateral line on blind side.

Interlinear scale rows

Frequencies

15 16 17 18 19

10 14 4 1 1

Fins: Dorsal with 101-110 (M = 107) rays, anal

with 80-89 (M = 84) rays, caudal 10 in 12 specimens (radiographs).

Vertebrae: 51-55 comprising 9 abdominal and

42-46 caudal elements in 12 specimens (radiographs).

Coloration: Upper side uniformly brownish,

opercular region rather blackish, lower whitish in

preserved specimens.

Size: Largest specimen examined, 230 (212 + 18)

mm, is from the Indian Ocean trawled by Anton

Bruun.

FIGURE 40.—Outline drawing of C. carpenteri (BMNH 1928.3.20.75) from Bay of Bengal.
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Seas of India to Persian Gulf;

trawled from a depth of 68-230 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus carpenteri, C. acutirostris, C. mar ley i, and C. suyeni are

four allopatric species that resemble each other in

the small, and closely placed eyes, wide mouth

cleft extending below or beyond the posterior border of fixed eye, and long pointed snout, all

specifications for living on a sandy or muddy bottom and plowing or burrowing into the substratum

for feeding. Cynoglossus carpenteri is closely related to C. acutirostris but can be separated from

it by its shorter snout (33.2 percent cf. 43.46 percent of head) and somewhat larger scales (15-19 cf.

18-20 interlinear rows).

TYPE SPECIMENS.—A specimen, ZSI 12434, 148

mm SL, from Orissa coast, coll. Marine Survey, is

here selected as the lectotype of C. carpenteri and a

specimen, ZSI 12433, 141.5 mm SL, collected along

with the lectotype as the paralectotype.

OTHER MATERIAL EXAMINED.—INDIA: 6 specimens, ZSI

12727, Bay of Bengal, 98-102 fms, coll. Marine Survey. 6,

ZSI 12728, Bay of Bengal, 98-102 fms, coll. Marine Survey. 6,

ZSI 13589-94, Bay of Bengal, coll. Marine Survey. 9, ZSI

F 647/1-656/1, off Calicut coast, 100 fms, coll. Marine

Survey. 6, ZSI F 1275/1-1280/1, Arabian Sea, 156-200 fms,

coll. Marine Survey. 3, BMNH 1890.7.31.10-12, Ganjam

coast, lat. 18°30'N, long. 84°36'E, coll. Marine Survey. 3,

BMNH 1890.11.28.27-29, Ganjam coast, lat. 18°30'N, long.

84°46'E, coll. Marine Survey. 1, BMNH 1928.3.20.74, Arabian

Sea, 156-200 fms, ex. Indian Museum. 3, BMNH 1928.3.20.

75-77, Bay of Bengal, 28-102 fms, ex. Indian Museum. 3,

BMNH 1928.3.20.78-80, off Calicut coast, 100 fms, coll.

Marine Survey. 53, SOSC Ref 23, Arabian Sea, lat. 24°13'N,

long. 65°52'E, 93 fms, coll. Anton Bruun. 1, SOSC Ref 23,

Arabian Sea, lat. 24°10'N, long. 65°54'E, 126 fms, coll.

Anton Bruun. 19, SOSC Ref 23, Arabian Sea, lat. 23°13'N,

long. 66°40'E, 100 fms, coll. Anton Bruun. 18, SOSC Ref

23, Arabian Sea, lat. 24°55'N, long. 61°10'E, 100 fms, coll.

Anton Bruun. 116, SOSC Ref 23, Arabian Sea, lat. 24°13'N,

long. 65°52'E, coll. Anton Bruun. 2, USNM 44418, Bay of

Bengal, coll. RIMS Investigator. OMAN: 4, ZSI F 1160/1-

63/1, Gulf of Oman, 230 fms, coll. Marine Survey. 3, BMNH

1901.1.30-32, Sea of Oman, 24°59'N, 60°58'E, 100 fms. coll.

Tounsend. 2, BMNH 1901.1.81.14-15, Gulf of Oman,

25°35'N, 57°47'E, 170 fms, coll. Tounsend. 1, BMNH

1939.5.24.1799, Gulf of Oman, 196 fms, coll. John Murray

Expd. PERSIA: 3, BMNH 1903.7.8.24-26, Persian Gulf,

15 fms, coll. Tounsend.

38. Cynoglossus acutirostris Norman

FIGURE 41; PLATE 17

Cynoglossus (Areliscus) acutirostris Norman, 1939:104, fig. 35

[type-locality: Gulf of Aden].

DESCRIPTION.—Based on 11 specimens, 182.0-

238.0 mm SL, including lectotype and paralectotypes of C. acutirostris.

Depth of body 26.29-30.53 (M = 28.40), length

of head 23.11-27.47 (M = 26.04) percent of standard length. Diameter of eye 6.14-9.71 (M = 7.95),

interorbital space rather narrow, 0.91-3.88 (M =

1.92) percent of length of head. Two nostrils on

ocular side, anterior nostril tubular in front of

fixed eye, posterior nostril simple, immediately in

front of interorbital space. Snout acutely pointed,

37.74-48.54 (M = 43.46) percent of head; rostral

hook rather long, reaches below anterior border

of fixed eye. Maxillary extending beyond fixed

eye; angle of mouth extending to below vertical

from posterior border of fixed eye or just beyond,

much nearer to branchial opening than to tip of

snout, tip of snout to angle of mouth 52.83-65.05

(M = 58.40), angle of mouth to branchial opening

35.79-45.28 (M = 39.90) percent of length of head.

Scales: Cycloid on ocular side with ctenoid scales

posteriorly; cycloid on blind side.

Lateral-Line System: Three lateral lines on ocular side, both dorsolateral and ventrolateral lines

slightly undulating and entering dorsal and anal

fins, respectively, at a very short distance from

caudal base; midlateral line with 84-92 (M = 87)

scales, 18-20 (M = 19) scales between them. No

lateral line on blind side.

Interlinear scale rows

Frequencies

18 19 20

5 4 2

Fins: Dorsal with 110 rays, anal with 94 rays,

caudal 10 in 1 specimen (radiograph).

Vertebrae: 58, comprising 9 abdominal and 49

caudal elements in 1 specimen (radiograph).

Coloration: Upper side uniformly brownish,

lower whitish in preserved specimens.

Size: Largest specimen examined, 254 (238 + 16)

mm, is one of the type specimens from Gulf of

Aden.

DISTRIBUTION.—Gulf of Aden; trawled from a

depth of about 120 fms.

DIAGNOSIS AND AFFINITIES.—The acutely pointed

long snout is unique among the species of the

genus Cynoglossus and serves as a ready diagnostic

character. In most other characters C. acutirostris is

very similar to C. carpenteri.

TYPE SPECIMENS.—A specimen, BMNH 1939.5.

24.1800, 238 mm SL, from Gulf of Aden, coll.
NUMBER 238 85

cm

a

FIGURE 41.—C. acutirostris, lectotype (BMNH 19395.24.800): a, outline

drawing of lateral view of ocular side; b, details of head of blind side.

John Murray Expd. is here selected as the lectotype of C. acutirostris and 9 specimens, BMNH

1939.5.24.1801-1809, 182-238 mm SL, collected

along with the lectotype and a specimen, USNM

109493, 190 mm SL, also collected along with the

lectotype are the paralectotypes.

39. Cynoglossus tnarleyi Regan

FIGURE 42; PLATE 18

Cynoglossus (Trulla) tnarleyi Regan, 1921b:418 [type-locality:

off Umvoti River, Natal].

Areliscus marleyi.—Bonde, 1922:25.—Barnard, 1925:414.—

Fowler, 1925:187.—Bonde, 1925:292.—Fowler, 1935:361.—

Barnard, 1937:41.—Smith, 1949:166.

DESCRIPTION.—Based on 2 specimens, 162.0-316.0

mm SL, including the holotype of C. marleyi.

Depth of body 21.60-25.63 (M = 23.62), length

of head 18.0-20.68 (M= 19.34) percent of standard

length. Diameter of eye 7.0-8.96 (M = 7.98), interorbital space 2.63-4.48 (M = 3.56) percent of

length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior

nostril simple, in anterior half of interorbital

space. Snout pointed, 38.81-45.61 (M = 42.41) percent of length of head; rostral hook rather long,

reaches below fixed eye. Maxillary extending to

below posterior border of fixed eye, or a little beyond it, angle of mouth extending to below vertical from posterior border of fixed eye, nearer to

branchial opening than to tip of snout; tip of snout

to angle of mouth 52.24-59.65 (M = 55.95), angle

of mouth to branchial opening 40.30-47.37 (M =

43.34) percent of length of head.
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

I cm

FIGURE 42.—Outline drawing of C. marleyi, holotype (BMNH 1921.3.1.21) from Natal.

Scales: Ctenoid on both sides.

Lateral-Line System: Three lateral lines on ocular side, midlateral line with 112-1 IS scales, 18-19

scales between middle and upper lateral lines. No

lateral line on blind side.

Interlinear scale rows 18 19

Frequencies 1 2

Fins: Dorsal with 126 rays, anal with 105 rays,

caudal 10 in 1 specimen (radiograph).

Vertebrae: 62, comprising 9 abdominal and 53

caudal elements in 1 specimen (radiograph).

Coloration: Upper side uniform brown, lower

whitish in preserved specimens.

Size: Largest specimen examined, 336 (316 + 20)

mm, is the holotype from Natal.

DISTRIBUTION.—Durban to Delagoa Bay; recorded from depths of 30-150 fms.

DIAGNOSIS AND AFFINITIES.—Cynoglossus marleyi,

which is restricted in its distribution to South

Africa, is related to C. carpenteri and is undoubtedly evolved from a carpenteri-lilne ancestor.

The ctenoid scales on the ocular side and the more

elongate snout (mean length 42.21 percent of head

cf. 33-32 percent) distinguish it from C. carpenteri.

TYPE SPECIMENS.—Holotype of C. marleyi,

BMNH 1921.3.1.21, 316 mm SL, from Natal, 130

fms, coll. Marley.

OTHER MATERIAL EXAMINED.—1 specimen, 160 mm SL, from

off Natal coast, lat. 29°27'S, long. 31°31'E, coll. Anton Bruun,

25.9.64.

40. Cynoglossus suyeni Fowler

PLATE 18

Cynoglossus xiphoideus [not Gunther].—Weber, 1913b:441.—

Weber and de Beaufort, 1929:193 [Timor Sea].

Cynoglossus suyeni Fowler, 1934a:347, fig. 101 [type-locality:

Mindora Is., Philippines].

Cynoglossus beauforti Chabanaud, 1951c:3, figs. 2, 3 [typelocality: Timor Sea].

DESCRIPTION.—Based on 20 specimens, 110.0-

257.0 mm SL, including the holotypes of C. suyeni

and C. beauforti.

Depth of body 19.29-25.25 (M = 22.59), length

of head 19.39-23.23 (M = 21.13) percent of standard length. Diameter of eye 6.67-12.28 (M = 9.10)

percent of length of head, interorbital space mostly

absent, or very narrow. Two nostrils on ocular side,

anterior nostril tubular in front of lower eye,

posterior nostril simple, in anterior part of interorbital space. Snout rather pointed, 38.46-46.34

(M = 42.97) percent of length of head; rostral

hook long, reaches below anterior border of fixed

eye. Maxillary extending well beyond fixed eye;

angle of mouth extending just behind vertical

from posterior border of fixed eye, nearer to branchial opening than to tip of snout; tip of snout to

angle of mouth 53.33-60.98 (M = 57.61), angle of

mouth to branchial opening 39.02-48.31 (M •»

44.05) percent of length of head.

Scales: Ctenoid on both sides.

Lateral-Line System: Three lateral lines on ocular side; midlateral line with 102-126 (M = 114)

scales, 19-22 (M = 20) scales between middle and

upper lateral lines. No lateral line on blind side.
NUMBER 238 87

Interlinear scale rows

Frequencies

19 20 21 22

4 9 3 1

Fins: Dorsal with 115-126 (M = 110) rays, anal

with 92-105 (M = 91) rays, caudal 10 in 9 specimens (radiographs).

Vertebrae: 57-61, comprising 9 abdominal and

48-52 caudal elements in 9 specimens (radiographs).

Coloration: Upper side uniformly brownish,

lower whitish in preserved specimens.

Size: Largest specimen examined, 275 (257 +18 )

mm, is from Philippine-Celebes area, trawled by

Albatross.

DISTRIBUTION.—From Philippines through Celebes to the Timor Sea; trawled from a depth of

173 fms.

DIAGNOSIS AND AFFINITIES.—Cynaglossus suyeni,

which is restricted to the deeper waters off Mindanao Island southward through the Celebes Sea

to the Timor Sea, is closely related to C. carpenteri

and has undoubtedly evolved from a carpenterilike ancestor. It can be readily separated from

C. carpenteri by the nature and position of its

eyes, which are scaly and contiguous. The ctenoid

scales on the ocular side and the longer snout

(mean length 42.97 percent of head cf. 33.2 percent)

further distinguish it from C. carpenteri. The scaly

covered contiguous eyes are considered a further

specialization for a burrowing mode of life.

NOTE ON SYNONYMY.—Cynoglossus beauforti

Chabanaud (1951c) was described based on a

specimen obtained from the Timor Sea by the

Siboga Expedition and included under C. xiphoideus by Max Weber (1931:441). Weber and de

Beaufort (1929:193) in giving the description of

C. xiphoideus (not Gunther) observed:

We made the above description after the specimen of the

Siboga Expedition from the Timor sea, mentioned by one of

us l.c, who stated already that there are some differences

between Cunther's type specimen and this one. In the type

the eyes are separated by more than one eye diameter and

the upper one is considerably in advance of the lower; also

the height is less, 4.2/3 times in length with caudal and the

head is shorter, 5]/z times in length with caudal. Therefore

we are not sure that our specimen really is the same as

C. xiphoideus and thought it therefore better to give a

description of this specimen only.

I have examined the holotype of C. beauforti

and compared it with the holotype and 13 other

specimens of C. suyeni in the National Museum

of Natural History, Smithsonian Institution, Washington, D.C. I have no hesitation regarding the

inclusion of it in the synonymy of C. suyeni.

TYPE SPECIMENS.—Holotype of C. suyeni, USNM

93086, 142 mm SL, from Mindanao Island, Philippines, coll. Albatross. Holotype of C. beauforti,

ZMA 108.001, 210 mm SL from Timor Sea, lat.

10°27'09"S, long. 123°28'7"E, coll. M. Weber,

Siboga Expd.

OTHER MATERIAL EXAMINED.—13 specimens, USNM 137941-

50, from Philippines to Celebes, coll. Albatross.

The heterolepis group

Two related species complexes of cynoglossus

can be assembled in what may be called the heterolepis group, heterolepis being closest to the evolutionary stem of the group. They are characterized

by a much elongated, pointed snout (except C.

semilaevis and C. macrophthalmus) with the angle

of mouth situated nearer to branchial opening

than to tip of snout, much smaller body scales, the

interlinear scale count being 15-25, very small eyes

with the interorbital space measuring about equal

to the diameter of the eye or even wider than one

eye diameter (C. heterolepis, C. gracilis, C. semilaevis, C. microlepis), and usually two nostrils on

ocular side and three lateral lines. No lateral line

on blind side.

The heterolepis group has its center of distribution in the Malay Archipelago and extends

through the South China Sea and Taiwan to Korea

and Japan; it is regarded as a very lately evolved

group of specialized species split off from the carpenteri complex.

The heterolepis complex

Cynoglossus heterolepis, C. feldmanni, C. wandersi, C. kapuasensis, C. semilaevis, C. abbreviatus,

C. gracilis, and C. microlepis form the heterolepis

complex. The characteristics of this complex are

almost the same as given for the group and it

differs from the macrophthalmus complex by

having both nostrils present.

41. Cynoglossus heterolepis Weber

FIGURE 43; PLATE 19

Cynoglossus heterolepis Weber, 1910:237 [type-locality:
88

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

b FIGURE 43.—C. heterolepis: a, outline drawing of holotype

(BMNH 1913.12.15.36) from Lorentz River, New Guinea; b,

details of the anal side of the holotype (female).

Lorentz R.]; 1913a:590 [Lorentz R., Bivak Is.].—Weber

and de Beaufort, 1929:186, figs. 52, 53 (Lorentz R.).—

Hardenberg, 1941:227 (Petpemboewe).—Munro, 1958:285.

DESCRIPTION.—Based on 4 specimens, 65-194 mm

SL, including the holotype of C. heterolepis.

Depth of body 16.44-23.45 <M = 18.89), length

of head 19.07-25.09 (M - 23.28) percent of standard length. Diameter of eye 3.23-5.97 (M = 5.08),

interorbital space 3.23-5.71 (M = 4.37) percent of

length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior

nostril simple, in the middle of interorbital space.

Snout obtusely pointed, 41.94-45.71 (M = 43.92)

percent of length of head; rostral hook rather

short, ends just in front of anterior nostril. Maxillary extending to a little behind fixed eye; angle of

mouth extending below vertical from posterior

border of fixed eye, nearer to branchial opening

than to tip of snout; tip of snout to angle of mouth

43.24-57.14 (M = 51.06), angle of mouth to

branchial opening 41.79-43.24 (M = 42.46) percent

of length of head.

Scales: Ctenoid on both sides, those on blind side

rather weakly serrated.

Lateral-Line System: Three lateral lines on

ocular side, midlateral line with 100-120 scales,

15-16 scales between middle and upper lateral line.

No lateral line on blind side.

Interlinear scale rows 15 16

Frequencies 2 2

Fins: Dorsal with 113-116 (M = 115) rays, anal

with 86-89 (M = 88) rays, caudal 10 in 4 specimens

(radiographs).

Vertebrae: 51-52, comprising 9 abdominal and

41-43 caudal elements in 4 specimens (radiographs).
NUMBER 238 89

Coloration: Upper side uniformly brownish with

or without darker spots, lower whitish in preserved

specimens.

Size: The largest specimen examined, 203 (194 +

9) mm, is the holotype from the Lorentz River, New

Guinea.

DISTRIBUTION.—New Guinea (Lorentz River) and

Australia (Northern Territory).

DIAGNOSIS AND AFFINITIES.—Though geographically widely separated, C. heterolepis is closely

related to C. feldmanni, found in the rivers of

Borneo, Sumatra, and Cambodia. It differs from

C. feldmanni chiefly in its slender body (mean depth

18.89 percent of SL, cf. 27.11 percent) and relatively large scales, the interlinear scale rows being

15-16 versus 17-18 in C. feldmanni.

TYPE SPECIMENS.—Holotype of C. heterolepis,

BMNH 1913.12.15.36, 194 mm SL, Lorentz River,

New Guinea, coll. M. Weber.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH 1937.3.

17.1, Upper Fly R., Papua, ex. American Museum. 4, USNM

174028, Australia, NT., coll. R. R. Miller, Arnhem Land

Expd.

42. Cynoglossus feldmanni (Sleeker)

FIGURE 44

Plagusia feldmanni Bleeker, 1853b:455 [type-locality: Borneo].

Arelia feldmanni.—Bleeker, 1859:184.

Cynoglossus feldmanni.—Bleeker, 1875:31, pi. 243: fig. 5, 184.

Cynoglossus (Areliscus) hardenbergi Norman, 1931.—422

[type-locality: Palembang, Sumatra].

Cynoglossus abentoni Stauch, 1966.—126 [type-locality: Baklau and Prek-Tasom, Cambodia].

DESCRIPTION.—Based on 2 specimens, 148.5 and

207.0 mm SL, including the holotype of C. hardenbergi.

Depth of body 26.94-27.29 (M = 27.11), length

of head 21.55-22.46 (M = 22) percent of standard

length. Diameter of eye 5.38-7.81 <M = 6.59), interorbital space 4.30-4.69 (M = 4.54) percent of

length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior

nostril simple, in anterior half of interorbital space.

Snout obtusely pointed, 39.78-40.62 percent of

length of head, rostral hook of moderate size,

reaches just below anterior border of fixed eye.

Maxillary extending to below posterior half or

posterior border of fixed eye; angle of mouth extending to below vertical from posterior half of

fixed eye, slightly nearer branchial opening than

tip of snout; tip of snout to angle of mouth 49.46-

51.46 (M = 50.46), angle of mouth to branchial

opening 46.88-49.46 (M = 48.17) percent of length

of head.

Scales: Ctenoid on both sides except those on the

head region of the blind side, where the scales are

either cycloid or very feebly ctenoid.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 81-84 scales, 17-18 scales

between middle and upper lateral lines. No lateral

line on blind side.

Interlinear scale rows 17 18

Frequencies 1 1

Fins: Dorsal with 98 rays, anal with 77 rays,

caudal 10 in 1 specimen (radiograph).

I cm

FIGURE 44. Outline drawing of C. feldmanni, holotype (BMNH 1862.6J.10) from Borneo.
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vertebrae: 50, comprising 9 abdominal and 41

caudal elements in 1 specimen (radiograph).

Coloration: Upper side brownish with irregular

darker spots, lower whitish in preserved specimens.

Size: Largest specimen examined, 231.5 (207 +

24.5) mm, is the type of C. hardenbergi from

Sumatra.

DISTRIBUTION.—Borneo, Sumatra, and Cambodia.

In rivers.

DIAGNOSIS AND AFFINITIES.—Cynoglossus feldmanni resembles both C. heterolepis and C. waandersi. The differences between this species and C.

heterolepis have been previously outlined. It is

distinguishable from C. waandersi by a lower interlinear scale count (17-18 cf. 24) and by the position

of the rectus of the mouth, which is nearer to the

branchial opening than to the tip of the snout

(rectus of the mouth in C. waandersi is much nearer

to the snout than to branchial opening).

NOTE ON SYNONYMY.—Bleeker (1853b:455) described P. feldmanni on the basis of a single specimen, 160 mm in TL from Pengaron, and characterized it as having three lateral lines on left side,

the upper and lower separated from the middle by

17 or 18 series of scales, and by the angle of the

mouth being nearer to gill opening than to extremity of snout. Norman (1931:422) described a similar

species C. (Areliscus) hardenbergi on the basis of a

single specimen, 233 mm in TL, from Palembaney,

Sumatra and related it to C. feldmanni, but differentiated it by its larger scales and longer and more

pointed rostral hook. He characterized the fish as

having three lateral lines on ocular side, the upper

and the middle separated by 16 or 17 series of scales.

I have examined the holotype of C. (Areliscus)

hardenbergi in the British Museum and found it

to conform well with the description and figure of

C. feldmanni, the differences noted by Norman

being considered here as attributable to intraspecific variation in the species.

Stauch (1966:126) described C. abentoni on the

basis of two specimens, 66 and 129 mm in TL, from

Ba-Klaut and Prek-Tasom, Cambodia, and characterized it as having three lateral lines on ocular

side, the upper and the middle being separated by

16 series of scales. Stauch, while describing the fish,

compared it with C. waandersi (Bleeker). Cynoglossus waandersi, however, has smaller scales, the

interorbital scale rows being 24, whereas in the

interlinear scale character and other proportional

and meristic characters C. abentoni is identical with

C. feldmanni. I have examined the holotype and

the paratype of C. abentoni in Paris and found

them identical with C. feldmanni.

TYPE SPECIMENS.—Among the Bleeker collection

in Leiden is a specimen, 145 mm SL, 162 mm TL,

cataloged as RMNH 6782 from Pengaran, Borneo,

coll. A. Feldmann, 1852 (pers. comm. Dr. M.

Boeseman) that appears to be the holotype of C.

feldmanni. Holotype of C. hardenbergi, BMNH

1931.4.23, 207 mm SL, Sumatra, coll. Hardenberg.

Holotype of C. abentoni, MNH P 65-466, 120 mm

SL, Ba Klaut, Patit Lac, coll. Aubenton, paratype,

MNH P 65-465, 61 mm SL, Prek Tasom, coll.

Aubenton.

OTHER MATERIAL EXAMINED.—SUMATRA: 1 specimen,

ZMA 108.004, Ojambi. BORNEO: I, BMNH 1862.6.3.10,

Borneo, coll. Bleeker.

43. Cynoglossus waandersi (Bleeker)

PLATE 19

Plagusia waandersii Bleeker, 1854a:98 [type-locality: confluence of rivers Lamatang and Enirn at Pelembang, eastern

Sumatra].

Arelia waandersii.—Bleeker, 1859:185.

Cynoglossus waandersi.—Bleeker, 1875:31, pi. 241: fig. 2.—

Weber and de Beaufort, 1929:189.

DESCRIPTION.—Based on a single specimen, 73.0

mm SL.

Depth of body 27.40 percent of standard length,

length of head 24.66 percent. Diameter of eye 11.11,

interorbital space 5.56 percent of length of head.

Two nostrils on ocular side, anterior nostril tubular, in front of lower eye, posterior nostril simple,

immediately in front of interorbital space. Snout

obtusely pointed, 41.67 percent of length of head;

rostral hook, rather short, ends in front of anterior

border of fixed eye. Maxillary extending to below

posterior half of fixed eye; angle of mouth extending below vertical from middle of fixed eye,

nearer to tip of snout than branchial opening; tip

of snout to angle of mouth 47.22, angle of mouth

to branchial opening 55.56 percent of length of

head.

Scales: Ctenoid on both sides.

Lateral-Line System: Three lateral lines on

ocular side, midlateral line with 80 scales, 24 scales

between middle and upper lateral lines. No lateral

line on blind side.
NUMBER 238 91

Fins: Dorsal with 89 rays, anal with 67 rays,

caudal 10 in 1 specimen (radiograph).

Vertebrae: 42, comprising 9 abdominal and 33

caudal elements (radiograph).

Coloration: Upper side yellowish brown, with

darker patches, lower whitish in preserved

specimens.

Size: The only specimen examined, 83 (73 + 10)

mm, is from Sumatra (reported up to 300 mm by

Weber and de Beaufort).

DISTRIBUTION.—Sumatra and Borneo, in rivers.

DIAGNOSIS AND AFFINITIES.—This species is more

closely allied to C. kapuasensis than to C. feldmanni

in having the more enlarged body and smaller

scales. It can, however, be separated from C.

kapuasensis by having the rictus of the mouth

nearer to the tip of the snout than to the branchial

opening and by its reduced eyes (diameter 11.11

percent in length of head cf. 7.41 percent).

From all the other members of the heterolepis

group, however, C. waandersi is distinct in its fewer

vertebrae (Figure 53, Graph 4).

TYPE SPECIMENS.—Bleeker's (1854a:98) description of P. waandersi was based on a single specimen,

145 mm in TL, from the confluence of the Lamatang and Enim rivers at Pelembang, Sumatra. In

his Atlas (1875-1876:31, pi. 241: fig. 2), he described

and illustrated a specimen measuring 145 mm in

TL. Among the Bleeker collection in Leiden is a

specimen, 129 mm SL, 145 mm in TL from the

confluence of Lamatang and Enim rivers, Prov.

Palembang, Sumatra, coll. P. S. van Bloemen

Waanders, 1854, cataloged as RMNH 6781 (pers.

comm. Dr. M. Boeseman). It is here considered to

be the holotype of C. waandersi.

OTHER MATERIAL EXAMINED.—1 specimen, USNM 171680,

Moesi R., Moeara, Sumatra, coll. A. Thieremann, 1929.

44. Cynoglossus kapuasensis Fowler

Cynoglossus kapuasensis Fowler, 1905:519 [type-locality:

Kapuas R., W Borneo].

Cynoglossus kapwasensis.—Seale, 1910:288 [misprint for C.

kapuasensis].

DESCRIPTION.—Based on a single specimen, the

holotype, 285.0 mm SL.

Depth of body 23.16, length of head 18.95 percent of standard length. Eyes small, diameter of

eye 7.41, interorbital space is equal to one eye diameter, 7.41 percent in length of head. Two nostrils

on ocular side, anterior nostril tubular, just in

front of vertical through front border of lower

eye, posterior nostril simple, in posterior half of

interorbital space. Snout rather obtusely pointed,

45.37 percent of head; rostral hook rather long,

reaches below fixed eye. Maxillary extending to

below posterior border of fixed eye; angle of mouth

extending to below vertical from posterior half of

fixed eye, nearer to branchial opening than to tip

of snout; tip of snout to angle of mouth 61.11, angle

of mouth branchial opening 50.0 percent of head.

Scales: Ctenoid on both sides, scales on anterior

third of body smaller than those on rest of body.

Lateral-Line System: Three lateral lines on

ocular side, dorsolateral line running backward and

entering dorsal fin along 5th ray, counted from the

rear, midlateral line with 126 scales, 25 scales between middle and upper lateral lines; ventrolateral

line runs backward and enters anal fin along 5th

ray, counted from the rear. No lateral line on blind

side.

Fins: Dorsal with 109 rays, anal with 85 rays,

caudal 10 in 1 specimen (radiograph).

Vertebrae: 50, comprising 9 abdominal and 41

caudal elements, in 1 specimen (radiograph).

Coloration: Upper side rather dark brown with

darker mottlings and blotches, lower rather brownish white in preserved specimens.

Size: The only specimen examined is 285.0 mm

in SL.

DISTRIBUTION.—Western Borneo (Kapuas River).

DIAGNOSIS AND AFFINITIES.—This species is closely

related to C. feldmanni and C. waandersi. It is distinguished from C. feldmanni by larger eyes (diameter 7.41 percent of head cf. 6.59 percent), wider

interorbital space (7.41 percent of head cf. 4.54

percent), and smaller scales, the interlinear scale

rows being much larger (25 cf. 17-18). From C.

waandersi it is separated by having the rictus of

the mouth situated nearer to the branchial opening

than to the tip of snout, by smaller eyes (diameter

of eye 7.41 percent of head cf. 11.11 percent), and

wider interorbital space (7.41 percent of head cf.

5.56 percent of head).

TYPE SPECIMEN.—Holotype of C. kapuasensis,

ANSP 72348, 285 mm SL, Kapuas R., W Borneo,

coll. Harrison and Hiller, 1898.
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

45. Cynoglossus semilaevis Giinther

FIGURE 45; PLATE 20

Cynoglossus semilaevis Gunther, 1873:379 [type-locality:

Chefoo, China]; 1898:261 [Nin-Tchouang].—Mori, 1927:183.

—Chu, 1931:95.—Wu, 1932:152 [Tsing-Wang-Tao, TsingTao, Tchefoo, Tcha-Pou, Wen Chou, and Amoy].

Areliscus semilaevis.—Jordan and Starks, 1906a:242.

Trulla semilaevis.—Fowler, 1934b:212.

Arelia rhomaleus Jordan and Starks, 1906b:526, fig. 5 [Port

Arthur, Manchuria].

Areliscus rhomaleus.—Jordan and Metz, 1913:63.—Reeves,

1927:14 [Swatow].—Sowerby, 1930:183.—Chu, 1931:95.

Cynoglossus roulei Wu, 1932:153 [type-locality: Amoy, China].

DESCRIPTION.—Based on 8 specimens, 134.0-530.0

mm SL, including the holotype of C. semilaevis and

syntypes of C roulei.

Depth of body 24.63-30.19 (M = 27.09), length

of head 20.89-26.79 (M = 23.93) percent of standard length. Upper eye not in advance of fixed eye,

diameter of eye 3.52-8.11 (M = 5.77), interorbital

space rather broad, 5.88-9.28 (M = 8.09) percent of

the length of head. Two nostrils on ocular side, anterior nostril tubular, in front of lower edge of

lower eye, posterior nostril simple, in the anterior

half of interorbital space. Snout rather rounded,

34.26-37.10 (M = 36.02) percent of head, rostral

hook short, extending to front of anterior nostril.

Maxillary extending to behind fixed eye; angle of

mouth extending below vertical from posterior half

of fixed eye, nearer to tip of snout than to branchial

opening; tip of snout to angle of mouth 44.33-50.0

(M = 46.69), angle of mouth to branchial opening

50.0-56.45 (M = 54.35) percent of length of head.

Scales: Ctenoid on ocular side, cycloid on blind

side.

Lateral-Line System: Three lateral lines on ocular

FIGURE 45.—C. semilaevis: a, outline drawing of specimen

(BMNH 1898558.9) from Liao-hu, north China; b, details

of anal region of blind side of holotype, male (BMNH

1873.953.16) from Chefoo.
NUMBER 238

93

side, midlateral line with 110-138 (M = 120) scales,

20-25 (M = 23) scales between middle and upper

lateral lines. No lateral line on blind side.

Interlinear scale rows

Frequencies

20 21 23 24 25

1 1 S 2 1

Fins: Dorsal with 124 rays, anal with 96 rays,

caudal 10 in 1 specimen (radiograph).

Vertebrae: 55, comprising 11 abdominal and 44

caudal elements in 1 specimen (radiograph).

Coloration: Upper side uniformly brown, lower

whitish in preserved specimens.

Size: Largest specimen examined, 566 (530 + 36)

mm, is from Shanghai.

DISTRIBUTION.—China.

DIAGNOSIS AND AFFINITIES.—This species is closely

related to C. abbreviatus but is readily separated

from it by the number of caudal fin rays (10), short

rounded snout (mean length 36.0 percent of head

cf. 37.52 percent), smaller eyes (mean diameter 5.77

percent of head cf. 7.57 percent), wider interorbital

space (8.09 percent of head cf. 6.80 percent), and

cycloid scales on the blind side of the body.

In the rounded short snout, 10 rays in the caudal

fin, and the position of the rictus of the mouth,

which is nearer to the snout than to the gill opening, C. semilaevis is considered a more generalized

species and could represent the ancestral condition

from which the more specialized species like C.

abbreviatus, C. gracilis, and C. microlepis evolved.

NOTE ON SYNONYMY.—Gunther (1873b:379) described C. semilaevis on the basis of a single specimen, 455 mm in TL, from Chefow, China and

characterized it as having a deeper body, three lateral lines on the ocular side, the middle and the

upper separated by 21 rows of scales, the scales between lateral lines smooth, between outer and vertical fins ctenoid, and cycloid on the blind side.

Arelia rhomaleus was described by Jordan and

Starks (1906b: 526) on the basis of one specimen

from Port Arthur, 380 mm in TL, and characterized

by having three lateral lines, the middle and the

upper being separated by 24 scale rows, ctenoid

scales on ocular side and cycloid on the blind side.

Jordan and Starks considered this species to be

closely related to C. abbreviatus but preferred to

retain it separately because of its increased number

of interlinear scale rows. From trigrammus (= abbreviatus) they differentiated this species by its

longer dorsal fin, larger eyes, and narrower interoribtal space. Fowler (1934b:213), however, confused this species with C. abbreviatus. Another

species C. roulei with almost identical characters as

C. rhomaleus was described by Wu (1932:153) from

China. I have examined a type specimen of all three

species and found them to be so similar in all

respects that there is no question but that they are

synonymous.

TYPE SPECIMENS.—BMNH 1873.9.23.16, 440.5

mm SL, holotype of C. semilaevis, Chefoo, coll.

Surinhoe. USNM 55635, 363 mm SL, holotype of A.

rhomaleus, Port Arthur, coll. J. F. Abbott. 2 specimens, 223 and 260 mm SL, BMNH 1860.7.28.48 and

BMNH 1924.12.15.64, types (? syntypes) of C. roulei,

Amoy, China, coll. Stevens (1860.7.28.48) and Light

(1924.12.15.64).

OTHER MATERIAL EXAMINED.—2 specimens, BMNH

1895.5.15-16, Shanghai, coll. Stylan. 1, BMNH 1896.258.9,

Liao-ho, N China, coll. Dr. W. Morrison. 1, BMNH

1930.7.28.13, Chekiang, China, coll. J. Chen. 3, BMNH

1939.1.17.56, Hong Kong. 1, (skeleton) MCZ 11323, Hong

Kong. 3, USNM 56395, Port Arthur, China, coll. J. F. Abbott.

1, USNM 137988, Kowloon, China, coll. Albatross.

46. Cynoglossus abbreviatus (Gray)

FIGURE 46; PLATE 20

Plagusia abbreviata Gray, 1835, pi. 94: fig. 3 [type-locality:

China].—Richardson, 1846:280 [coasts of China, Canton].

Cynoglossus abbreviatus Gunther, 1862:494 [Amoy]; 1873a:244

[Shanghai].—Bleeker, 1873:136.—Peters, 1880:923 [Ningpo].

—Seale, 1914:78 [Hong Kong].—Fowler and Bean, 1920:321

[Soochow].—Evermann and Shaw, 1927:113 [Hangchow,

Nanking, Ningpo].—Tchang, 1929:37, fig. 43 [Yangtze R.].—

Fowler, 1930:615 [Hong Kong].—Wu, 1932:157 [Tchou-San,

Tchefoo, Tsingtao].—Ochiai, 1963:95, pi. 21.—Chu,

1963:544, fig. 441.—Shen, 1967:208 [Kowloon].

Areliscus abbreviatus.—Jordan and Metz, 1913:62 [Fusan].—

Reeves, 1927:14 [Shanghai, Ningpo, Swatow].—Wu, 1929:71.

fig. 57 [Amoy].—Chu, 1931:%.—Herre and Myers, 1931:252

[Canton].—Matsubara, 1955:1286.

Trulla abbreviata.—Fowler, 1934b:213, fig. 29 [China, Hong

Kong, Canton, Amoy, Soochow, Ningpo, Hongchow, Nanking, Shanghai, Yangtze R.].

Cynoglossus trigrammus Gunther, 1862:494, [type-locality:

Chinese seas].—Bleeker, 1865b:56 [Amoy]; 1873:131.—Martens, 1876:399 [Shanghai].—Sauvage, 1881:107 [Swatow].—

Rutter, 1897:89 [compiled].—Wu, 1932:160 [Swatow].—Wu

and Wang, 1933:303 [Amoy].—Ochiai, 1959:211; 1963:92,

pi. 20.—Chu, 1963:542, fig. 409.—Shen, 1967:210 [Kowloon].

Areliscus trigrammus.—Reeves, 1927:14.—Oshima, 1927:202

[Taihoku].—Chu, 1931:95.—Herre. 1932:433 [Canton].—

Matsubara. 1955:1286.

Trulla trigrammus.—Fowler, 1934b:214.
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Areliscus purpureomaculatus Regan, 1905a:26 [type-locality:

Inland Sea of Japan].—Jordan and Starks, 1906a:242.—

Jordan, Tanaka, and Snyder, 1913:336.—Jordan and Hubbs,

1925:303.—Wu, 1932:156 [Tsing-Tao, Chefoo].—Kamohara,

1952:86 [Tosa].—Mori, 1952:184 [Pusan].

Cynoglossus xiphoideus [not Giinther].—Wu, 1932:158.

Areliscus gracilis [not Giinther].—Chen and Weng, 1965:99

[Tungkong].

DESCRIPTION.—Based on 19 specimens, 96.0-285.0

mm SL, including the types of C. trigrammus and

the holotype of C. purpureomaculatus.

Depth of body 22.58-27.78 (M = 24.98), length of

head 18.16-22.22 (M = 20.22) percent of standard

length. Diameter of eye 5.56-9.52 (M = 7.57), interorbital space 4.35-9.47 (M = 6.80) percent of length

of head. Two nostrils on ocular side, anterior nostril

tubular, in front of lower eye, posterior nostril

simple, in anterior half of interorbital space. Snout

obtusely pointed, 35.24-40.71 (M = 37.52) percent

of length of head, rostral hook short, ends in front

of anterior nostril. Maxillary extending to below

posterior half or rarely to posterior border of fixed

eye; angle of mouth extending below vertical from

middle or posterior half of fixed eye, slightly nearer

to tip of snout than to branchial opening; tip of

snout to angle of mouth 46.23-50.0 (M = 47.89),

angle of mouth to branchial opening 47.62-55.79

(M = 52.44) percent of head.

Scales: Rather small, ctenoid on both sides, those

on blind side weakly serrated.

Lateral-Line System: Three lateral lines on ocular

side, dorsolateral line running backward and entering dorsal fin posteriorly, midlateral line with 118—

136 (M = 125) scales, 18-23 (M = 21) scales between middle and upper lateral line, ventrolateral

line runs backward and enters anal fin posteriorly.

No lateral line on blind side.

Interlinear scale

Frequencies

rows 18

3

19

1

20

4

21

6

22

4

23

1

Fins: Dorsal with 128-135 (M = 130) rays, anal

with 100-108 (M = 104) rays, caudal 8 (9) in 12

specimens (radiographs).

Vertebrae: 60-64, comprising 10-11 abdominal

and 46-53 caudal elements in 12 specimens (radiographs).

Coloration: Upper side brownish with or without

dark blotches, lower whitish in preserved specimens.

Size: Largest specimen examined, 295.0 mm SL,

is from Hong Kong.

DISTRIBUTION.—South China Sea (Amoy, Canton,

Kowloon, Hong Kong, and Shanton) through Taiwan, Shanghai, Tunghai, and Po-hai to Korea and

Japan.

DIAGNOSIS AND AFFINITIES.—Cynoglossus abbreviatus is closely related to C. gracilis; it is distinguished by its deeper body (mean depth of body

24.98 percent of SL, cf. 20.78 percent) and larger

eyes (mean diameter 7.57 percent of head cf. 40.25

percent).

NOTE ON SYNONYMY.—Gray's naming of P. abbreviatus was based on Hardwicke's collection of

illustrations. The illustration that is the iconotype

of the species was issued in 1834 as part of one

group among several that were later combined in

two volumes without text (Gray, 1834). Giinther

(1862:496) gave the description of the species and

characterized it as having three lateral lines, the

middle and the upper separated by 19 series of

scales, the height of the body being contained three

and three-fourths in the total length, and the length

of head four times and two-thirds. This description

agrees well with Gray's illustration of the species.

Giinther mentioned the type of the species "adult:

km

FIGURE 46.—Outline drawing of C. abbreviatus (BMNH 1924.12.15.62) from Amoy.
NUMBER 238 95

stuffed. China. Presented by J. R. Reeves Esq." I

have examined two stuffed specimens, 220 and 315

mm SL, of C. abbreviatus from Reeves' collections,

which in most probability formed the basis for

Hardwicke's illustration of the species.

Cynoglossus trigrammus, described on the basis of

two specimens, 182.5 and 275 mm SL, from China

is a synonym of P. abbreviatus. The differences

noted, such as 21 series of scales separating the middle and upper lateral line, the height of the body

four times and two-thirds and the length of the

head five times and one-third are all attributable to

intraspecifk variation. Most subsequent workers,

however, treated C. trigrammus as a species distinct

from C. abbreviatus (Fowler, 1934b:214), their

differentiation of the two species being mainly

based on the depth of the body in TL. They considered forms with depth of three and three-fourths

to four as C. abbreviatus, and those above four to

four and one-half as C. trigrammus. Fowler further

considered T. abbreviata as a species with ctenoid

scales on the ocular side and cycloid on the blind

(Fowler, 1934b:214) and T. trigrammus with ctenoid on both sides. I have examined the types of

C. trigrammus and compared them with a number

of specimens labeled as C. abbreviatus from China

in the British Museum and found them to be similar in proportional and meristic characters. The

nature of the body scales as has been pointed out

earlier cannot be depended upon for species differentiation, as the ctenoid scales tend to become

weaker in their ctenoid nature as the fish grows. In

bigger specimens the scales on the blind side appear

more or less cycloid, especially on the anterior half

of the body. Obviously, Fowler's (1934b) inclusion

of Arelia rhomaleus in the synonymy of C. abbreviatus was based on the cycloid nature of scales of

the blind side of the fish, and A. rhomaleus is here

placed under C. semilaevis. Ochiai (1963) synonymized C. purpureomaculatus Regan with trigrammus. Cynoglossus purpureomaculatus was described

from Japan on the basis of a single specimen, 202

mm SL, which I have examined and found to agree

well with C. abbreviatus. Regan did not compare

his species with any other but indicated an interlinear scale row of 19, which falls within the range

of variation of this character in C. abbreviatus.

TYPE SPECIMENS.—Of the two specimens from

China on which Gunther based his description of C.

trigrammus, a specimen, BMNH 1856.9.19.1215,

182.5 mm SL, from the Hassler collection is selected here as the lectotype of C. trigrammus and

the other specimen, BMNH 1851.12.27.169, 155 mm

SL, as the paralectotype. BMNH 1905.6.6.248, 202

mm SL, holotype of C. purpureomaculatus, Japan,

coll. Gordon Smith.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH

1863.2.23.28. Amoy, China, coll. R. Surinhoe. 1, BMNH

1873.7.30, 60, Shanghai, coll. R. Surinhoe. 4,

BMNH 1924.12.15.58-61, Foochow, China. 1, BMNH

1924.12.15.62, Amoy, coll. Light. 1, BMNH 1924.12.15.63.

Amoy, China, coll. Light. 2, BMNH 1924.12.15.65, Foochow,

China, coll. Light. 2, BMNH 1930.7.28.10.11, Paihow, Canton, coll. J. Chen. 1, BMNH 1931.6.23.7, Pearl R., Canton,

coll. Hoffman. 2, BMNH 1939.1.17.57-58, Hong Kong, coll.

Herklot. 1, BMNH 1939.1.12.59, Hong Kong, coll. Harklots.

1, BMNH 1939.3.23, 100, Hong Kong, coll. Harklots. 2,

stuffed, China, coll. Reeves. 2, USNM 37762, Korea, coll.

J. B. Bernadou. 1, USNM 84006, China, coll. N. Gist Gee.

1, USNM 86028, China, coll. Sowerby. 2, USNM 87050,

China, coll. Sowerby. USNM 87051, China, coll. Sowerby.

3, USNM 130447, China, coll. Sowerby. USNM 130482,

China, coll. Sowerby. 1, USNM (without reg. no.). Pearl

River mouth, Hong Kong, coll. Hong Kong Fisheries Dept.

1, ANSP 52908, Hong Kong coll. F. H. Fowler. 2, MCZ

11155-11156, China. 1, UMMZ 167447. China, coll. J. J. Tu.

47. Cynoglossus gracilis Gunther

FICURE 47; PLATE 21

Cynoglossus gracilis Gunther, 1873a:244 [type-locality: Shanghai]; 1898:261 [New Chwang].—Mori, 1927:184.—Wu.

1932:160 [Yangtze R.].—Fowler, 1934b:214.—Tchang.

1955:304, fig. 189.—Chu, 1963:543, fig. 410.—Shen. 1967:209

[Kowloon].

Areliscus gracilis.—Reeves, 1927:14 [Chefoo].—Chu, 1931:96.—

Matsubara, 1955:1286.

Trulla gracilis.—Fowler, 1934b:214.

Cynoglossus microps Steindachner, 1898:510 [type-locality:

Hong Kong].—Wu, 1932:158.

Areliscus hollandi Jordan and Metz, 1913:62, pi. 9: fig. 3

[type-locality: Fusan, Korea].

Cynoglossus pellegrini Wu, 1932:159 [type-locality: Hainan].

Areliscus trigrammus [not Gunther].—Chen and Weng,

1965:98, fig. 68 [Quemoy].

DESCRIPTION.—Based on 14 specimens (84-228

mm SL), including holotype and paratypes of C.

gracilis.

Depth of body 19.08-23.20 (M = 20.78), length of

head 16.92-23.18 (M = 20.42) percent of standard

length. Upper eye not in advance of lower, diameter

of eye 4.21-7.84 (M = 5.86), interorbital space 5.00-

8.16 (M = 6.27) percent of length of head. Two

nostrils on ocular side, anterior nostril tubular, in
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 47.—Outline drawing of C. gracilis, holotype (BMNH 1873.7.30.57), from Shanghai.

front of lower eye, posterior nostril simple, in anterior half of interorbital space, or at the middle

of it. Snout obtusely pointed, 35.29-46.58 (M =

40.25) percent of length of head, rostral hook of

moderate size, usually extending up to anterior

nostril but sometimes reaching below anterior border of fixed eye. Maxillary extending to behind

fixed eye; angle of mouth extending to below vertical from posterior border of fixed eye, nearer to

branchial opening than to tip of snout; tip of snout

to angle of mouth 47.73-57.53 (M = 52.14), angle

of mouth to branchial opening 47.37-52.73 (M =

48.96) percent of length of head.

Scales: Large or moderate, ctenoid on both sides,

those on blind side, particularly in large specimens,

very weakly serrated with a much reduced number

of ctenii.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 122-138 (M = 135) scales,

19-22 (M = 21) scales between middle and upper

lateral lines. No lateral line on blind side.

Interlinear scale rows 19 20 21 22

Frequencies 2 2 9 2

Fins: Dorsal with 133-137 (M = 135) rays, anal

with 104-108 (M = 106) rays, caudal 8 in 10 specimens (radiographs).

Vertebrae: 62-64, comprising 11 or 12 abdominal

and 51-55 caudal elements in 10 specimens (radiographs).

Coloration: Upper side uniformly light brown,

lower whitish in preserved specimens.

Size: Largest specimen examined, 278.0 mm, is

from liao-hu, N China.

DISTRIBUTION.—From South China Sea through

Taiwan, Tung-hai, and Chefoo to Pusan in Korea.

DIAGNOSIS AND AFFINITIES.—Cynoglossus gracilis

is closely related to C. abbreviatus. It is distinguished from C. abbreviatus by its elongate body

(mean depth 20.78 percent of cf. 24.98 percent),

smaller eyes (5.86 percent of head cf. 7.57 percent),

and longer snout (mean length 40.25 percent of

head cf. 37.52 percent).

NOTE ON SYNONYMY.—Gunther (1873a:244) described C. gracilis on the basis of one specimen,

225 mm in TL, and several young ones from Shanghai and characterized it as a species having three

lateral lines, the upper and the middle being separated by 21 scales, extremely small eyes, and

elongated body, the depth of body being five times

in total length. The descriptions of C. microps and

C. pellegrini do not indicate any significant difference from C. gracilis, and in both these species the

height of the body is contained more than four

times in the TL. Jordan and Metz (1913:62) described A. hollandi from Fusan, Korea on the

basis of a single specimen and from the description

and illustration of the fish, its identity with C.

gracilis is unmistakable.

TYPE SPECIMENS.—BMNH 1873.7.30.57, 206 mm

SL, holotype of C. gracilis, Shanghai, coll. Swinhoe.

2, BMNH 1873.7.30.58a, 59b, 87. and 79.5 mm SL,

paratypes of C. gracilis, obtained along with the

holotype. NHV 43.798, 227 mm SL, 241 mm TL,

from Yangtse-Kiang, selected as lectotype of C.

microps, NHV 43.799, 190 mm SL, 203 mm TL,

collected along with lectotypes is a paralectotype of

C. microps.
NUMBER 238 97

OTHER MATERIAL EXAMINED.—2 specimens, BMNH

1873.9.23.20-21, Chefoo, coll. Swinhoe. 1, BMNH 1898.2.28.10,

Liao-hu, N China, coll. Dr. W. Morrison. 4, BMNH

1924.12.15.54-57, Foochow, China, coll. Light. 1, BMNH

1930.7.28-12, Chekiang, China, coll. J. Chen. 1, USNM 86466,

China, coll. Sowerby. 3, USNM 130547, China, coll. Sowerby. 3, USNM 130521, Shanghai, China. 3, USNM 131071,

China, coll. Dale and Jony, 1881. 1, USNM 148414, Shanghai, coll. D. C. Janson. 1, USNM 11329, Hong Kong. 1,

NHV 43820, Shanghai. 1, NHV 43822, Canton. 1, ANSP

52907, Hong Kong, coll. H. W. Fowler.

48. Cynoglossus microlepis (Sleeker)

FIGURE 48; PLATE 21

Plagusia microlepis Bleeker, 1951a:413 [type-locality: Borneo];

1852:31.

Arelia microlepis.—Bleeker, 1859:184.

Cynoglossus microlepis.—Gunther, 1862:495.—Bleeker, 1875:32,

pi. 243: fig. 3.—Vaillant, 1893:101.—Weber and de Beaufort, 1929:190—Smith, 1933:83; 1945:442.—Suvatti, 1950:

327.—Punpoka, 1964:65.

Cynoglossus xiphoideus Gunther, 1862:495 [type-locality:

Siam].—Bleeker, 1865a:33.—Kner, 1867:294.—Hora, 1923a:

182.—Smith, 1933:83; 1945:441.—Suvatti, 1950:329.—Chabanaud, 1951c: 1.

Cynoglossus (Arelia) solum Sauvage, 1878:92 [type-locality:

Me Kong River].

Arelia solum Sauvage, 1883:151.

Cynoglossus feldmanni [not Bleeker].—Weber and de Beaufort [in part], 1929:192.

Cynoglossus trigrammus [not Gunther].—Suvatti, 1950:329.—

Punpoka, 1964:75 [Gulf of Thailand].

DESCRIPTION.—Based on 4 specimens, 142.0-231.0

mmSL.

Depth of body 20.07-22.51 (M = 20.84), length of

head 19.05-20.35 (M = 20.21) percent of standard

length. Migratory eye is much in advance of fixed

eye, diameter of eye 5.00-6.67 (M = 5.50), interorbital space rather broad 6.67-8.51 (M = 7.54)

percent in length of head. Two nostrils on ocular

side, anterior nostril tubular, in front of fixed eye,

posterior nostril simple, in posterior half of interorbital space. Snout obtusely pointed, 40.0-45.0

(M = 42.82) percent of length of head, rostral hook

rather long, reaches below middle or posterior half

km

FIGURE 48.—C. microlepis (BMNH 1859.7.132) from Thailand: a, outline drawing of lateral

view of ocular side; b, nature of scales along base of dorsal fin on ocular side; c, same on

blind side.
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of fixed eye. Maxillary extending to below posterior

border of fixed eye or a little behind; angle of

mouth extending to below vertical from posterior

half of fixed eye, nearer to branchial opening than

to tip of snout; tip of snout to angle of mouth

54.26-55.0 (M = 54.82), angle of mouth to branchial

opening 40.0-48.94 (M = 44.11) percent of length

of head.

Scales: Small, ctenoid on both sides, those on

blind side weakly serrated.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 116-138 scales, 21-22

scales between middle and upper lateral lines. No

lateral line on blind side.

Interlinear scale rows 21 22

Frequencies 5 2

Fins: Dorsal with 117-119 rays, anal with 92-96

rays, caudal 8 in 4 specimens (radiographs).

Vertebrae: 55-56, comprising 9 abdominal and

46-47 caudal elements in 4 specimens (radiographs).

Coloration: Upper side reddish brown with dark

spots, lower whitish in preserved specimens.

Size: Largest specimen examined, 304 (270 + 34)

mm, is from Sarawak, Borneo.

DISTRIBUTION.—Thailand, Cambodia, Vietnam,

Borneo, and Sumatra. In strictly fresh water.

DIAGNOSIS AND AFFINITIES.—Cynoglossus microlepis is closely related to C. gracilis in most morphometric and meristic characters but can be readily

distinguished by the position of its migratory eye,

which is placed much in advance of the fixed eye.

The much smaller eyes (mean diameter 5.50 percent

of head cf. 7.57 percent), broader interorbital space

(mean width 7.54 percent of head cf. 6.80 percent),

and longer snout (mean length 42.82 percent of

head cf. 37.52 percent) further distinguish the species from C. gracilis. Cynoglossus microlepis is considered more specialized toward a burrowing mode

of life than C. abbreviatus or C. gracilis.

NOTE ON SYNONYMY.—Bleeker (1851a:413) described P. microlepis on the basis of a single specimen, 94 mm in TL, from rivers of Bandjermassing

in Borneo, and characterized it as having three

lateral lines on ocular side, one on blind side, much

elongated body, the height of the body being five

times in TL, smaller eyes, and 150 scale rows along

the midlateral line. Unfortunately, neither Bleeker

nor Giinther (1862:495) gave the number of scale

rows that separate the upper and the middle lateral

lines in this species. This resulted in confusion as to

the correct identity of the species, causing the creation of several new ones. Cynoglossus xiphoideus

was described on the basis of two specimens, 150

and 229 mm in TL, from Thailand and characterized by three lateral lines on ocular side, the upper

and middle being separated by 20 or 21 scale rows,

one on the blind side, the interorbital space being

more than that of the orbit and the upper eye considerably in advance of the lower. Many workers

considered this species distinct from C. microlepis,

though in the excellent illustration of C. microlepis

in Bleeker's Atlas (pi. 243: fig. 3), 21-22 interlinear

scale rows and the position of the migratory eye in

advance of the fixed eye are clearly indicated. Smith

(1945:441) considered the two species distinct; the

principal point of differences noted was a single

distinct lateral line on the blind side in the case of

C. microlepis and the larger number of scales in the

medium lateral line as compared with C. xiphoideus. Comparison of a large number of specimens

from Thailand does not substantiate Smith's conclusions. The lateral line on the blind side is totally

absent in all the specimens examined, though a

somewhat lateral line-like streak is visible on the

blind side, which, of course, is not pored. The type

specimens of C. xiphoideus, in the British Museum,

that were examined agree well with Bleeker's description and illustration of P. microlepis, and since

this species has priority over C. xiphoideus, I have

used the name C. microlepis for the species.

Cynoglossus solum described by Sauvage (1878:92)

from the Mekong River and characterized as having

three lateral lines on the colored side, one on the

blind side, and 166 scales in the midlateral line

was synonymized by Smith (1945:442) and I concur

with him.

TYPE SPECIMENS.—Of the two type specimens of

C. xiphoideus in the British Museum, a specimen,

BMNH 1859.7.1.52, 191 mm in SL, is selected here

as the lectotype and the other specimen, BMNH

1859.7.1.53, 137.5 mm SL, as the paralectotype of

C. xiphoideus.

Among the Bleeker collection in Leiden is a

specimen, cataloged as RMNH 6784, 171 mm SL,

193 mm TL, from Bandjermassin, SE Borneo, coll.

J. Wolff (pers. comm., Dr. M. Boeseman), and it is

here considered as the holotype of C. microlepis. A.

second specimen, RMNH 17876, 325 mm SL, 360
NUMBER 238

mm TL, is the figured specimen indicated as such

by Chabanaud.

OTHER MATERIAL EXAMINED.—1 specimen, BMNH

1876.3.20.4, Laos, coll. Paris Museum. 5. BMNH 1898.4.2.130-

134, Menam R., ex. Siamese Museum. 2, BMNH

1906.10.29.39-40, Sarawak, coll. S. L. Hora. 1, BMNH

1928.5.22.2, Bangkok, Thailand, coll. H. M. Smith. 1,

ZMA 108.004, Indonesia, coll. Moolenburgh. 2, ANSP 89534.

Gulf of Thailand, coll. R. M. de Schauensee. 1. USNM

103319, Menam Chau Phya, Thailand. 1, UMMZ 181238,

Cambodia, coll. J. Bardoch. 1, ZSI F 19608, Montaburi,

1

Thailand, coll. H. M. Smith.

The macrophthalmus complex

This species is like the members of the heterolepis complex except that only one (posterior)

nostril is present on the ocular side. The center of

distribution is in Queensland and probably descended from the same ancestral stock from which

the heterolepis complex was derived.

49. Cynoglossus macrophthalmus Norman

PLATE 21

Cynoglossus macrophthalmus Norman, 1926:306, fig. 15 [typelocality: Southern Queensland, 20 miles off Bustard Head

light].

DESCRIPTION.—Based on 2 specimens, 73.0-251.0

mm SL, including the holotype of C. macrophthalmus.

Depth of body 27.40-28.97 percent of standard

length, length of head 23.84-24.66 percent. Diameter of eye 11.11-11.35, interorbital space 4.84-8.83

percent of length of head. Anterior nostril on ocular

99

side tubular, in front of lower eye, posterior nostril absent. Snout rounded, 35.06-38.89 percent of

length of head, rostral hook long, reaches below

posterior border of fixed eye or just beyond. Maxillary extending to just behind fixed eye; angle of

mouth extending to below vertical from posterior

border of fixed eye, slightly nearer to branchial

opening than to tip of snout; tip of snout to angle

of mouth 50.08-55.56, angle of mouth to branchial

opening 47.22-48.41 percent of length of head.

Scales: Small, weakly ctenoid on both sides.

Lateral-Line System: Three lateral lines on ocular

side, midlateral line with 100-103 scales, 17-18

scales between middle and upper lateral lines. No

lateral line on blind side.

Fins: Dorsal with 110-111 rays, anal with 90-91

rays, caudal 10 in 2 specimens (radiographs).

Vertebrae: 53, comprising 9 abdominal and 44

caudal elements in 2 specimens (radiographs).

Coloration: Upper side brownish, lower whitish

in preserved specimens.

Size: Largest specimen examined, 270 (251 + 19)

mm, is the holotype from southern Queensland.

DISTRIBUTION.—Queensland; recorded from a

depth of 20 fms.

DIAGNOSIS AND AFFINITIES.—Apparently related to

the C. heterolepis complex but differing in the

lack of a posterior nostril and possessing a rounded

snout and a long rostral hook reaching to below

posterior border of fixed eye. In its elongated body

and reduced eyes it resembles C. waandersi.

TYPE SPECIMEN.—Holotype of C. macrophthalmus, AMS E 1978, 251 mm SL, 20 miles off Bustard

Head light, southern Queensland.

OTHER MATERIAL EXAMINED.—1 specimen, AMS

1A 6460, from Lindeman Island, Queensland.
Literature Cited

•Not seen by author

Alcock, A.

1889. Natural History Notes from H. M. Indian Marine

Survey Steamer "Investigator," Commander Alfred

Carpenter, Commanding, No. 10: list of the Pleuronectidae Obtained in the Bay of Bengal in 1888

and 1889 with Descriptions of New and Rare Species. Journal of the Asiatic Society of Bengal,

58(2):279-295, plates 16-18.

1890a. Natural History Notes from H. M. Indian Marine

Survey Steamer, "Investigator," Commander Hoskyn,

Hoskyn, R. N., Commanding, No. 16: On the

Bathybial Fishes Collected in the Bay of Bengal

during the Season 1889-90. Annals and Magazine of

Natural History, series 6, 6:197-222.

1890b. Natural History Notes from H. M. Indian Marine

Survey Steamer, "Investigator," Commander Hyskyn,

R. N., Commanding, No. 20: On Some Undescribed

Shore Fishes from the Bay of Bengal. Annals and

Magazine of Natural History, series 6,6:425-443.

1896. Natural History Notes from H. M. Indian Marine

Survey Steamer, "Investigator," Series II, No. 23: A

Supplementary List of the Marine Fishes of India,

with Descriptions of Two New Genera and Eight

New Species. Journal of the Asiatic Society of Bengal, 65:301-333.

1898. Fishes. Part V in Illustrations of the Zoology of the

Royal Indian Marine Survey Ship, "Investigator."

Plates XVIII-XXIV.

1899. A Descriptive Catalogue of the Indian Deep-Sea

Fishes in the Indian Museum, being a Revised Account of the Deep-Sea Fishes Collected by the Royal

Indian Marine Survey Ship "Investigator." Pages 1-

11+ 1-211 + index. Calcutta: Government Press.

Barnard, K. H.

1925. A Monograph of the Marine Fishes of South Africa.

Annals of the South African Museum, 21:1-1065.

1937. Further Notes on South African Marine Fishes. Annals of the South African Museum, 32(2):41-67.

Ben-Tuvia, A.

1953. Mediterranean Fishes of Israel. Bulletin of the Sea

Fisheries Research Station (Haifa), 8:1-40, 20 figures.

1963. Systematics and Ecology of Indo-Pacific Fishes Recently Established in the Eastern Mediterranean.

International Congress of Zoology, 16:115. [Abstract.]

1966. Red Sea Fishes Recently Found in the Mediterranean. Copeia, 1966(2):254-275.

Bleeker, P.

1849. Bijdrage tot de kennis der ichthyologische fauna van

het eiland, Bali, met beschrijving van eenige nieuwe

species. Verhandelingen van het Bataviaasch genootschap van Kunsten en Wetenschappen, 22:1-11.

1851a. Over eenige nieuwe soorten van Pleuronectoiden

van den Indischen Archipel. Natuurkundig tijdschrift voor Nederlandsch-Indie, 1:401-416.

1851b. Bijdrage tot de kennis der ichthyologische fauna

van Riouw. Natuurkundig tijdschrift voor Nederlandsch-Indie, 2:469-497.

1852. Bijdrage tot de kennis der Pleuronectoidei van den

Soenda-Molukschen Archipel. Verhandelingen van

het Bataviaasch genootschap van Kunsten en Wetenschappen, 24:1-32.

1853a. Nalezingen op de ichthyologische fauna van Bengalen en Hindostan. Verhandelingen van het Bataviaasch genootschap van kunsten en wetenschappen,

25:1-164,6 plates.

1853b. Zevende bijdrage tot de kennis der ichthyologische

fauna van Borneo: Zoetwatervischen van Sambas,

Pontianak en Pengaron. Natuurkundig tijdschrift

voor Nederlandsch-Indie, 5:427-462.

1853c. Nieuwe tientallen diagnostische beschrijvingen van

nieuwe of Weinig bekende vischsoorten van Sumatra.

Natuurkundig tijdschrift voor Nederlandsch-Indie,

5:495-534.

1854a. Overzicht der ichthyologische fauna Van Sumatra,

met beschrijving van eenige nieuwe soorten. Natuurkundig tijdschrift voor Nederlandsch-Indie, 7:49-108.

1854b. Specierum piscium javanensium novarum vel minus

cognitarum diagnoses adumbrate. Natuurkundig

tijdschrift voor Nederlandsch-Indie, 7:415-448.

1858a. Twaalfde bijdrage tot de kennis der vischifauna van

Borneo: Visschen Van Sinkawang. Ada Societatis

Scientiarum Indo-Neerlandicae, 5:1—10.

1859a. Enumeratio specierum piscium hucusque in Archipelago Indico observatarum, adjectis habitation ibus

citationibusque ubi descriptiones earum recentiores

reperiuntur, nee non speciebus Musei Bleekeriani

Bengalensibus, Japonids, Capensibus, Tasmanicisque.

Ada Societatis Scientiarum Indo-Neerlandicae,

6:1-276, 36 plates.

1860a. Achtste bijdrage tot de kennis vischfauna van

Sumatra. Visschen Van Benkoelen, Priaman, Tandjong, Palembang en Djambi. Ada Societatis Scientiarum Indo-Neerlandicae, 8:1-88.

1960b. Dertiende bijdrage tot de kennis der vischfauna van

Borneo. Acta Societatis Scientiarum Indo-Neerlandicae, 8:1-64.

1865a. Nouvelle notice sur la faune ichthyologique de Siam.

Nederlandsch Tijdschrift voor de Dierkunde, 2:33-

37.

100
NUMBER 238 101

1865b. Notice sur les poissons envoye's de Chine au Mus6e

de Leide par M. G. Schlegel. Nederlandsch Tijdschrift voor de Dierkunde, 2:55-62.

1873. Mlmoire Sur la fauna ichthyologique de Chine.

Nederlandsch Tijdschrift voor de Dierkunde, 4:113—

154.

1875. Atlas Ichthyologique des Indes Orientates Neerlandaises. Volume 6(1866-1872), pages 1-170, plates

232-278.

Bonde, C. von

1922. The Heterosomata (Flat Fishes) Collected by the

S.S. "Pickle." Report of Fisheries and Marine Biological Survey, Union of South Africa, 2(Special

Report, l):l-29, plates 1-6.

1925. A Collection of Heterosomata from Portugese East

Africa. Transactions of the Royal Society of South

Africa, 12:285-294.

Botros, T. A.

1971. Fishes of the Red Sea. Oceanography and Marine

Biology, an Annual Review, 9:221-348.

Boulenger, G. A.

1901. Notes on the Classification of Teleostean Fishes, IV:

On the Systematic Position of the Pleuronectidae.

Annals and Magazine of Natural History, 7(8):261-

271.

1902. Flat Fishes of the Cape Colony. Marine Investigations in South Africa, 1:1-10.

•Buttikofer, J.

1890. Reisebilder aus Liberia. 2 volumes.

Cadenat, J.

1960. Poissons de mer ouest-africaine observes du Senegal

au Camoroun et plus specialement au large des

cotes de Sierra Leone et du Ghana. Bulletin de

I'Institut Francaise d'Afrique Noire, 22 A: 1358-1430,

6 plates.

Cantor, T.

1850. Catalogue of Malayan Fishes. Journal of the Royal

Asiatic Society of Bengal, 18[1849]:i-xii + 983-1442,

14 plates.

•Capello, M. D. B.

1872. Journal de Sciencias mathematicas physicas et

naturaes publicado sob as auspicias da Academia

real das Sciencias de Lisboa.

Castelnau, F. L. de L. de

1861. Memoire sur les Poissons de I' Afrique australe.

Volume 7, i-vii + 1-78 pages.

Chabanaud, P.

1931. Sur divers poissons soleiformes de la region IndoPacifique. Bulletin de la SocUti Zoologique de

France, 56:291-305.

1933. Contribution a 1' osteologie comparative des poissons, principalement des teleosteens heterosomes.

Bulletin de la Sociite1 Zoologique de France, 58:140-

168, 1 plate, 21 figures.

1934a. Poissons recueillis dans le lac Timsah (isthme de

Suez) per M. le Professeur A. Gruvel, en 1933. Bulletin du Museum National d' Histoire Naturelle de

Paris, (2)6:156-160.

1934b. Heterogenite des tel£ost£ens dyssymetriques. Bulletin

de la Socie'ti Zoologique de France, 59:275-284, 14

figures.

1936. Le neurocrane osseua des teleosteens dyssyme'trique

apres la metamorphose. Annales de I'Institut Oceanographique, 16:223-297, 112 figures.

1937. Les teleosteens dyssymetriques du Mokattam Interieur de Tourah. Me'moires Institut d' £gypte,

32:1-121, 4 plates, 19 figures.

1939. Catalogue syst&natique et chorologique des teHost£ens dyssymetriques du Globe. Bulletin de I'

Institut Oceanographique, 763:1-31 + list of addenda and corrigenda.

1940. Contribution a la morphologie des Cynoglossidae

(Teleostei Pleuronectoidea Soleiformes). Bulletin du

Museum National d' Histoire Naturelle de Paris,

(2)12:182-191,6 figures.

1947a. Notules ichthyologiques, XXXII: Description d" un

nouveau Cynoglossus de la Cdte du Senegal. Bulletin

Museum National d' Histoire Naturelle de Paris,

(2)19:440-443.

1947b. Notules ichthyologiques, XXXIII: Definition d'un

genre inedit, appartment a la famille des Cynoglossidae. Bulletin du Museum National d' Histoire

Naturelle de Paris, series 2, 19:443.

1947c. Description d'un nouveau Cynoglossus de 1' Inde.

Annals and Magazine of Natural History, series 11,

14:813-815.

1949a. Revision des Cynoglossidae (S. Str.) de 1' Atlantique

Oriental. Bulletin du Museum National d' Histoire

Naturelle de Paris, series 2, 21:60-66.

1949b. Revision des Cynoglossidae (S. Str.) de 1' Atlantique

Oriental (Suite). Bulletin du Museum National

d' Histoire Naturelle de Paris, series 2, 21:202-209.

1949c. Revision des Cynoglossidae (S. Str.) de Atlantique

Oriental (Suite & fin). Bulletin du Museum National

d' Histoire Naturelle de Paris, series 2, 21:347-353.

1949d. Les Cynoglossus de 1' Atlantique. Bulletin du Museum National d' Histoire Naturelle de Paris, series

2.21:516-521.

1949e. Description d'un nouveau Cynoglossus de la Mer

Rouge. Bulletin de la Societe Zoologique de France,

74:145-148.

1950. Notules ichthyologique, XLII-XLVI. Bulletin du

Museum National d' Histoire Naturelle de Paris,

series 2, 22:336-340, 1 figure.

1951a. Definition et nomenclature des morphes pleurogrammiques des Cynoglossidae: Revision de quatre

especes du genre Cynoglossus. Bulletin du Museum

National d' Histoire Naturelle de Paris, series 2.

22:713-716.

1951b. Definition et nomenclature des morphes pleurogrammique des cynoglossidae: Revision de quatre especes

du genre Cynoglossus. Bulletin du Museum National

d' Histoire Naturelle de Paris, series 2, 23:77-81.

1951c. Sur deux Cynoglossus de la collection ichthyologique

der Zoologisch Museum, Amsterdam. Beaufortia,

5:1^4, 3 figures.

1951d. Sur divers Cynoglossus de la region Indo-Padfique.

Annals and Magazine of Natural History, series 12,

4:262-273.
102

1954. Notules ichthyologiques, Notule 47: Presence inddite

d'un Cynoglossus dans la Mediterrane orientate.

Bulletin du Museum National d' Histoire Naturelle

de Paris, series 2, 26:465-466.

1955a. Rectifications afferentes a la nomenclature et 4 la

syst&natique des pleuronectiformes du Sous—ordre

des Soleoidei. Bulletin du Museum National d' Histoire Naturelle de Paris, series 2, 27:447-452.

1955b. Sur la repartition circumafricaine de diverses especes

de la famille des Soleidae. Comptes Rendus Hebdomadaires des Seances et Memoires de la Societi

de Biogeographie, 281:99-106.

1956. Sur divers pleuronectiformes apparement au naturhistorisches museum de vienne: Nouvelle description

de Cynoglossus canariensis. Osterreichs Zoologische

Zeitschrift, 5(1955):403-406.

1965. Notules ichthyologique 50: Modification avec l'age

et l'ecaillure zenithala, phenomene paraissant particulier a certains teleosteens soleidei du genre

Cynoglossus. Bulletin du Museum d'Histoire Naturelle de Paris, series 2, 36:756-760.

Chabanaud, P., and T. Monod

1926. Les poissons de Port-Etienne: Contribution a la

faune ichthyologique de a region du Cap Blanc

(Mauritanie Franchise). Bulletin du Comiti d'ttudes

Historiques et Scientifiques de I'Afrique Occidentale

Frangaise, 6:225-287, 2 plates, 33 figures.

Chen, J. T. F.

1951. Monograph of the Fishes of Taiwan (in Chinese).

Quarterly Journal of the Taiwan Museum, 4:110—

163.

•1956. Synopsis of the Vertebrates of Taiwan.

Chen, J. T. F., and H. T. C. Weng

1965. A Review of the Flat Fishes of Taiwan. Biological

Bulletin, (Tunghai University), 25-27(Ichthyology

Series, 5): 1-103, 72 figures.

Chu, Y. T.

1931. Index Piscium Sinensium. Biological Bulletin of St.

John's University (Shanghai), 1:1-290.

1963. Fishes of Tung-hai. Pages 510-544.

Cuvier, G.

1836. Le regne animal. 3rd edition, volume 4, 391 pages,

120 plates.

Davies, A. M.

1934. Tertiary Faunas, a Text Book for Oilfield Palaeontologists and Students of Geology. Volume 2, i-x +

1-252 pages, 22 figures. London: Thomas Murby &

Co.

Day, F.

1867. On Some New or Imperfectly Known Fishes of

Madras. Proceedings of the Zoological Society of

London, pages 558-565.

1869. Remarks on Some of the Fishes in the Calcutta Museum. Proceedings of the Zoological Society of London, pages 511-527,548-560, 611-614.

1873. On Some New Fishes of India. Journal of the Linnean Society, 11:524-529.

1877. Fishes of India. Part 3, pages 369-552, plates 79-

133. London.

1889. Fishes. In Fauna of British India, including Ceylon

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and Burma. Volume 2, pages 1-XIV + 1-506, 177

figures. London: Taylor & Francis.

De Silva, P. H. D. H.

1956. The Flat Fishes (Heterosomata) of Ceylon with a

List of the Specimens in the Colombo Museum.

Ceylon Journal of Science, Section C, Fisheries, 7

(2): 183-200.

Dum^ril, A. H. A.

1858. Reptiles et poissons de l'Afrique occidentale: Etude

precede de considerations generates sur leur distribution geographique. Archives du Museum National d' Histoire Naturelle, 10:137-268.

Duncker, G.

1904. Die Fische der malayischen Halbinsel. Mitteilungen

aus dem Naturhistorischen Museum, 21:133-207.

Ekman, S.

1953. Zoogeography of the Sea. 417 pages. London: Sidgurick & Jackson Ltd.

Evans, H. M.

1937. A Comparative Study of the Brains in Pleuronectidae. Proceedings of the Royal Society, series B,

122:308-43, 30 figures.

Everman, R. W., and T. Shaw

1927. Fishes from Eastern China, with Descriptions of

New Species. Proceedings of the California Academy

of Sciences, 16:97-122.

Fowler, H. W.

1904. A Collection of Fishes from Sumatra (with bibliography). Journal of the Academy of Natural Sciences

of Philadelphia, 2(12):495-560, 21 plates.

1905. Some Fishes from Borneo. Proceedings of the Academy of Natural Sciences of Philadelphia, 57:455-

523.

1918a. The Fishes of the United States Eclipse Expedition

to West Africa. Proceedings of the United States

National Museum, 56:195-292.

1918b. A List of Philippine Fishes. Copeia, 58:62-65.

1925. Fishes from Natal, Zululand, and Portuguese East

Africa. Proceedings of the Academy of Natural Sciences of Philadelphia, 77:187-268, 4 figures.

1930. Notes on Japanese and Chinese Fishes. Proceedings

of the Academy of Natural Sciences of Philadelphia,

81:589-616, 2 figures.

1934a. Descriptions of New Fishes Obtained in 1907 to

1910 Chiefly in the Philippine Islands and Adjacent

Seas. Proceedings of the Academy of Natural Sciences

of Philadelphia (1933), 85:233-367, 117 figures.

1934b. A Synopsis of the Fishes of China, Part V (continued): The Cods, Opahs, Flounders, Soles, John

Dories, Berycoids, Pipe Fishes, Silversides, Mullets,

Barracudas, and Thread Fishes. Hong Kong Naturalist, 5(3):21O-225. 9 figures.

1935. South African Fishes Received from Mr. H. W.

Bell-Marley in 1935. Proceedings of the Academy

of Natural Sciences of Philadelphia, 87:361-408, 39

figures.

1936. The Marine Fishes of West Africa, Based on the

Collection of the American Museum Congo Expedition, 1909-1915. Bulletin of the American Museum

of Natural History, 70(1): 1-1493,567 figures.
NUMBER 238 103

1938. A List of the Fishes Known from Malaya. 268 pages.

Singapore: Government Printing Press.

1956. Fishes of the Red Sea and Southern Arabia, 240

pages, 115 figures. Jerusalem: Weizmann Science

Press of Israel.

Fowler, H. W., and B. A. Bean

1920. A Small Collection of Fishes from Soochow, China

with Descriptions of Two New Species. Proceedings

of the United States National Museum, 58:307-321.

1922. Fishes from Formosa and the Philippine Islands.

Proceedings of the United States National Museum,

62:1-73. 4 figures.

Fox, H. M.

1926. Zoological Results of the Cambridge Expedition to

the Suez Canal, 1924. Transactions of the Zoological

Society of London, 22:1-64.

Frizzel, D. L., and J. P. Dante

1965. Otoliths of Some Early Coenozoic Fishes of the

Gulf Coast. Journal of Paleontology, 39(4):687-718,

3 plates.

Gilbert, C. H.

1889. A Preliminary Report on the Fishes Collected by the

Steamer "Albatros" on the Pacific Coast of North

America during the year 1889, with Description

of Twelve New Genera and Ninety-Two New Species. Proceedings of the United States National Museum, 13:49-126.

Gilchrist, J. D. F.

1905. Description of New South African Fishes. Marine

Investigations in South Africa, 3:1-16.

1908. Descriptions of Fifteen New South African Fishes

with Notes on Other Species. Marine Investigations

in South Africa, 4:143-171.

Gilchrist, J. D. F., and W. W. Thompson

1917. A Catalogue of the Sea Fishes Recorded from

Natal. Annals of Durban Museum, 1(4):291-431.

Ginsburg, J.

1951. Western Atlantic Tongue-Fishes with Descriptions

of Six New Species. Zoologica, 36(3): 185-201.

Goode, G. B., and T. H. Bean

1885. Descriptions of New Fishes Obtained by the United

States Fish Commission, Mainly from Deep Water

of the Atlantic and Gulf Coasts. Proceedings of the

United States National Museum, 8:589-605.

1886. Report on the Results of Dredging, under the

Supervision of Alexander Agassiz in the Gulf of

Mexico (1877-78) and the Caribbean Sea (1878-79)

and along the Atlantic Coast of the United States

by the U.S. Coast Survey Steamer "Blake," XXXVI:

Oceanic Ichthyology. Bulletin of the Museum of

Comparative Zoology at Harvard College, 12:1—

xxxV +1-553, 123 plates (417 figures).

Gray, John Edward

1835. Illustrations of Indian Zoology, Chiefly Selected

from the Collections of Major General Hardwicke.

Volume 2, 102 plates. London: Adolphus Richter

& Co.; Parbury, Allen 8c Co.

Gruvel, A., and P. Chabanaud

1937. Missions A. Gruvel dans le canal de Suez, II:

Poissons. Memoires Institut d' Egypte, 35:1-30, 29

figures.

Gunther, A.

1861. Catalogue of the Fishes in the British Museum.

Volume 3, 586 pages.

1862. Catalogue of the Fishes in the British Museum.

Volume 4, 504 pages.

1873a. Report on Collections of Fishes from China. Annals

and Magazine of Natural History, series 4, 12:239-

250.

1873b. On a Collection of Fishes from Chefoo, North

China. Annals and Magazine of Natural History,

series 4, 12:377-380.

1878. Notes on a Collection of Japanese Sea-Fishes. Annals

and Magazine of Natural History, series 5, 1:485-487.

1880. Report on the Shore Fishes. Pages 1-82 in part 6 in

volume 1 of Zoology in Reports . . . the Voyage of

HM.S. "Challenger" 32 plates.

1898. Report on a Collection of Fishes from Newchwang,

North China. Annals and Magazine of Natural

History, 7(l):257-263, plate XIII.

Hamilton-Buchanan, F.

1822. An Account of the Fishes Found in the River

Ganges and Its Branches. 405 pages, 39 plates.

Edinburgh: Archibald Constable.

Hardenberg, J. D. F.

1941. Fishes of New Guinea. Treubia, 18:217-231, 4

figures.

Herre, A. W.

1932. Fishes from Kwangtung Province and Hainan

Island, China. Lingnan Science Journal (Canton),

11:423-443, 1 figure.

1933. A Check List of Fishes from Sandakan, British

North Borneo. Journal of the Pan-Pacific Research

Institute, 8:2-5.

1934. Notes on Fishes in the Zoological Museum of

Stanford University, 1: The Fishes of the Herre

Philippine Expedition of 1931. Pages 1-106.

1941. A List of Fishes Known from the Andaman

Islands. Memoirs of the Indian Museum, 13:331-403.

1953. Check List of Philippine Fishes. U.S. Fish and Wildlife Service, Research Report (United States Department of Interior), 20:1-977.

Herre, A. W., and G. S. Myers

1931. Fishes of the South-eastern China and Hainan.

Lingnan Science Journal (Canton), 10:233-254.

Honma, Y.

1952. A list of the Fishes Collected in the Province of

Echigo, including Sado Island (in Japanese). Japanese Journal of Ichthyology, 2:220-229.

Hora, S. L.

1923a. On a Collection of Fish from Siam. Journal of the

Natural History Society of Siam, 6:143-184, plates

X-XII.

1923b. Fauna of the Chilka Lake. Memoirs of the Indian

Museum, 5:739-769.

1929. An Aid to the Study of Hamilton-Buchanan's

"Gangetic Fishes." Memoirs of the Indian Museum,

9:169-192, 10 plates.
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hubbs, C. L.

1915. Flounders and Soles from Japan, Collected by the

United States Bureau of Fisheries Steamer, "Albatross" in 1966. Proceedings of the United States

National Museum, 48:449-496, plates XXV-XXXII.

1945. Phylogenetic Position of the Citharidae, a Family of

Fiat-Fishes. Miscellaneous Publicaitons of the Museum of Zoology, University of Michigan, 93:1-38.

Jenkins, J. T.

1910a. Report on the Fishes Taken by the Bengal Fisheries Steamer "Golden Crown." Memoirs of the

Indian Museum, 3:1-31.

1910b. Notes on Fish from India and Persia, with Descriptions of New Species. Records of the Indian Museum, 5:123-140, 1 plate.

Jerdon, T. C.

1853. Ichthyological Gleanings in Madras. Madras Journal

of Literature and Science, 17(39[1851]): 128-151.

Johnstone, J.

1904. Report of the Marine Fishes Collected by Professor

Herdman, at Ceylon in 1902. Report of the Government of Ceylon on the Pearl Oyster Fisheries of the

Gulf of Mannar, part 2 (supplemental report),

15:201-222, 2 plates.

Jordan, D. S., and B. W. Evermann

1903. Notes on a Collection of Fishes from the Island

of Formosa. Proceedings of the United States National Museum, 25:315-368, 29 figures.

Jordan, D. S., and C. L. Hubbs

1925. Record of Fishes Obtained by David Starr Jordan

in Japan, 1922. Memoirs of the Carnegie Museum,

10(2):93-346, plates V-XII.

Jordan, D. S., and C. W. Metz

1913. A Catalogue of the Fishes Known from the Waters

of Korea. Memoirs of the Carnegie Museum, 6:1-65.

10 plates.

Jordan, D. S., and R. £. Richardson

1908. Fishes from Islands of the Philippine Archipelago.

Bulletin of the United States Bureau of Fisheries,

27:233-287.

1909. A Catalogue of the Fishes of the Island of Formosa

or Taiwan, Based on the Collections of Dr. Hans

Sauter. Memoirs of the Carnegie Museum, 4:159-204.

Jordan, D. S., and A. V. Seale

1907. Fishes of the Islands of Luzon and Panny. Bulletin

of the Bureau of Fisheries, 26:1-48.

Jordan, D. S., and J. O. Snyder

1900. A List of Fishes Collected in Japan by Keinosuke

Otaki, and by the United States Steamer "Albatross"

with Descriptions of Fourteen New Species. Proceedings of the United States National Museum,

23:335-380. plates 19, 20.

1901. A Preliminary Check List of the Fishes of Japan.

Annotationes Zoologicae japonenses, 3:31-159.

Jordan, D. S., and £. C. Starks

1906a. A Review of the Flounders and Soles of Japan.

Proceedings of the United States National Museum,

31:161-246.

1906b. Notes on a Collection of Fishes from Port Arthur,

Manchuria, Obtained by James Francis Albott.

Proceedings of the United States National Museum,

31:515-526.

Jordan, D. S., S. Tanaka, and J. O. Snyder

1913. A Catalogue of the Fishes of Japan. Journal of the

College of Science, Imperial University of Tokyo,

33(l):300-400.

Jordan, D. S., and W. F. Thompson

1914. Record of the Fishes Obtained in Japan in 1911.

Memoirs of the Carnegie Museum, 6:205-213.

Kamohara, T.

1934. Supplementary Notes on the Fishes Collected in the

Vicinity of Kochi Shi. Zoological Magazine (Tokyo),

46:457-463.

1938. On the Offshore Bottom-Fishes of Province Tosa,

Shikoku, Japan. 86 pages.

1952. Revised Descriptions of the Offshore Bottom-Fishes

of Province Tosa, Shikoku, Japan. Research Report

of the Kochi University, 3:1-122, 100 figures.

Kaup, J. J.

1858. Uebersicht der Plagusinae der funften subfamilie

der Pleuronectidae. Archiv fur Naturgeschichte,

24(1): 105-110.

Klunzinger, C. B.

1871. Synopsis der Fische des Rothen Meeres, II: Theil.

Verhandlunger der Zoologisch-botanischen Gesselschaft in Wien, 21:441-668.

1880. Die Von Mullersche Sammlung australischer Fische

in Stuttgart. Sitzungsberichte der Akademie der

Wissenschaften in Wien, 80(l):325-430.

Kner, R.

1867. Fische. Pages 275-433 in part 5 of Zoologischer

Theil of Reise der Ostereichischen Fregatte Novara

um die Erde in den Jahren 1857, 1858, 1859. 5

plates. Wien.

Kosswig, C.

1955. Zoogeography of the Near East. Systematic Zoology,

4(2):49-96.

Kuroda, N.

1931. list of the Fishes Found in the Waters Near

Shizuura, Province Suruga. Amoeba, 3:85-127.

1951. A Nominal List with Distribution of the Fish of

Suruga Bay, inclusive of the Fresh-Water Species

Found Near the Coast. Japanese Journal of Ichthyology, 1:314-338, 376-394.

1954. Additions to the List of Fishes of Suruga Bay.

Japanese Journal of Ichthyology, 3:64-67, 3 figures.

Kuronuma, K.

1939. The Miscellaneous Notes on the Flat Fishes Preserved at the Kominato Marine Biological Station,

Chiba Prefecture. Suisan-Kenkyusi, 34:83-86.

1961. A Check List of Fishes of Vietnam. 66 pages. Division of Agriculture and Natural Resources: United

States Operations Mission to Vietnam.

Kyle, H. M.

1913. Flatfishes (Heterosomata). Volume 2 in Report on

the Danish Oceanographical Expeditions, 1908-1910,

to the Mediterranean. 150 pages, 4 plates.

1921. The Asymetry, Metamorphosis, and Origin of Flat

Fishes. Philosophical Transactions of the Royal

Society, (B) 211:75-129.
NUMBER 238 105

Lacepede, B. G. £.

1802. Histoire Naturelle des Poissons, 4:l-XLIV + l-728,

16 plates.

Madeay. W.

1884. Supplement to the Descriptive Catalogue of Fishes

of Australia. Proceedings of the Linnean Society of

New South Wales, 9:2-64.

Marshall, N. B.

1952a. The Manihine Expedition to the Gulf of Aquaba,

1948-49, IX: Fishes. Bulletin of the British Museum

(Natural History), Zoology, l(8):221-252, 5 figures.

1952b. Recent Biological Investigations in the Red Sea.

Endeavour, 11:137-142.

Marshall. T. C.

1964. Fishes of the Great Barrier Reef and Coastal Waters

of Queensland. 566 pages, 136 plates. Sydney:

Angus and Robertson.

• Martens, E.

1876. Die preussische Expedition nach Ost'Asien, Zoologische Abtheilung. 2 volumes. Berlin.

Mauubara, K.

1955. Fish Morphology and Hierarchy. Parts 1 and 2,

pages 1-1605, 135 plates (461 figures), 267 textfigures. Tokyo.

Matthew, W. D.

1915. Climate and Evolution. Annals of the New York

Academy of Sciences, 24:171-318.

Menon, A. G. K.

1971. On the Scientific Identity of the Malabar Sole.

Journal of the Marine Biological Association of

India, 11(1,2) (1969):311-315.

•Mori, T.

1927. Handlist of Manchurian Vertebrates.

1952. Check List of the Fishes of Korea. Memoirs of the

Hyogo University of Agriculture, Biological Series,

1:1-228.

Munro, I. S. R.

1955. The Marine and Fresh Water Fishes of Ceylon.

Pages 1-XVI+1-349. 56 plates, Sydney: Halstead

Press.

1958. The Fishes of the New Guinea Region: A Check

List of the Fishes of New Guinea Incorporating

Records of Species Collected by the Fisheries Survey

Vessel "Fairwind" during the Years 1948 to 1950.

Fisheries Bulletin of the Department of Agriculture,

Stock and Fisheries (Papua and New Guinea), 1:97—

369. 1 figure.

Myers, G. S.

1952. The Nature of Systematic Biology and of Species

Description. Systematic Zoology, 1(3): 106-111.

Nielsen, J. G.

1961. Heterosomata (Pisces). Galathea Report, 4:219-226,

1 plate, 3 figures.

1964. Heterosomata (Pisces) Collected by Dr. Th.

Mortensen off South Africa. Videnskabelige Meddelelser fra dansk naturhistorisk Forening, 127:127-

134, 1 plate, 1 figure.

Nijssen. H.

1966. The Occurrence of Cynoglossus browni Chabanaud,

1949, in the North Sea. Beaufortia, 13:87-90.

Norman, J. R.

1925. Two New Fishes from China. Annals and Magazine

of Natural History, (9)16:270.

1926. A Report on the Fiat-Fishes (Heterosomata) Collected by the F.I.S. "Endeavour," with a Synopsis

of the Fiat-Fishes of Australia and a Revision of

the Subfamily Rhombosolinae. Biological Results,

"Endeavour," 5(5):219-3O8, 15 figures.

1928. The Fiat-Fishes (Heterosomata) of India, with a

List of the Specimens in the Indian Museum.

Records of the Indian Museum, 30(2): 173-215, 4

plates, 30 figures.

1931. Two New Fiat-Fishes from the Indo-Australian

Archipelago, with a Synopsis of the Species of the

Genera Poecilopsetta and Nematops. Treubia, IS:

421-426, 2 figures.

1934. A Systematic Monograph of the Fiat-Fishes (Heterosomata.): Psettodidae, Bothidae, Pleuronectidae. 456

pages, 318 figures. London: British Museum (Natural

History).

1939. Fishes. Number 1 in volume 7 of Scientific Reports,

John Murray Expedition. 116 pages, 41 figures.

Ochiai, A.

1959. Morphology, Taxonomy, and Ecology of the Soleoid

Fishes Found in Japan. 234 pages, 70 figures (in

Japanese). Tokyo.

1963. Fauna Japonica Soleina. 114 pages, 24 plates, 54

figures. Tokyo: Chiyoda Printing & Publishing Co.

Ltd.

1966. Studies on the Comparative Morphology and

Ecology of the Japanese Soles. Special Report of

Misaki Marine Biological Institute (Kyoto University), 3:1-97.

Ogilby. J. D.

1910. On New or Insufficiently Described Fishes. Proceedings of the Royal Society of Queensland, 23:1-55.

Okada, Y., and K. Matsubara

1938. Keys to the Fishes and Fish-like Animals of Japan,

including Kuril Islands, South Sakhalin, Bonirs

Islands, Ryukyu Islands, Korea and Formosa. 584

pages, 113 plates. Tokyo. [In Japanese.]

Oshima, M.

1927. list of Flounders and Soles Found in the Waters

of Formosa, with Descriptions of Hitherto Unrecorded Species. Japanese Journal of Zoology,

1:177-204.

Osorio. B.

1894. Estudes ichthyologieas aoerca da fauna das dominios

portuguezes na Africa. Journal de Sciencias mathematicas, physicas et naturaes publicado sob as

auspicias da Academic real das Sciencias de Lisboa,

series 2. 3:128-140.

1898. Da distribuicao geographica des Peixes e Crustacea*

Colhidas nas passessoes portuguezas d" Africa occidentale existentes no Museo national de Lisboa.

Journal de Sciencias mathematicas, physicas et

naturaes publicado sob as auspicias de Academia

real das sciencia de Lisboa, series 2, 5:185-202.

•1909. Contribuicao para o conhedmento da fauna bathy-
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pelagica visinha das costas de Portugal. Memoirs

Museum of Bocage, volume 1.

Otaki, K.

1897. Identification of the Fiat-Fishes Found in Japan.

Suisan Chiosa Hokoku, 6:17-26. [In Japanese.]

Parker, G. H.

1903. The Optic Chiasma in Teleosts and Its Bearing on

the Asymmetry of the Heterosomata (Flat Fishes).

Bulletin of the Museum of Comparative Zoology at

Harvard College, 40:221-242, 1 plate.

Pellegrin, J.

1905. Mission de pecheries de la Cote Occidentale d'

Afrique: Poissons. Actes de la Societe linneenne de

Bordeaux, 60:17-57.

1913. Poissons marins de Guinee, de la Cote d' Ivoire, du

Dahomeyet du Congo: Mission de M. Gruvel.

Bulletin de la Societe Zoologique de France, 38:151—

158.

1914. Missions Gruvel sur la Cote Occidentale d" Afrique

(1905-1912): Poissons. Annales de V Institut Oceanographique, 6(4): 1-100.

Pellegrin, J., and P. Chevey

1940. Poissons nouveaux ou rares de Cochinchine description de deux especes et de deux varietis. Bulletin

de la Societe Zoologique de France, 65:153-158, 2

figures.

Peters, W.

1880. A List of the Fishes from Ningpo Sent by the

Chinese Government to the Berlin Fish Exhibition.

Monatsberichte der Konigliche Akademie der Wissenschaften zw Berlin, 1880:921-927.

Piechocki, R.

1961. Wirbeltiere. Volume 1 of Makroskopische Praparationstechnik: Leitfaden fur das sammeln, Praparieren und Konservieren. 428 pages. Leipzig.

Pradhan, M. J.

1964. A Preliminary Account of the Fiat-Fishes (Heterosomata) Found along the Bombay Coast. Journal

of the Bombay Natural History Society, 61(2):456-

458.

Punpoka, S.

1964. A Review of the Fiat-Fishes (Pleuronectiformes =

Heterosomata) of the Gulf of Thailand and Its

Tributaries in Thailand. Bulletin of Fisheries Research (Kasetsart University), 1:1-86, 32 figures.

Reeves, C. D.

1927. A Catalogue of the Fishes of North-Eastern China

and Korea. Journal of the Pan-Pacific Research

Institution, 2:1-16.

Regan, C. T.

1905a. On a Collection of Fishes from Inland Sea of

Japan. Annals and Magazine of Natural History,

(7)15:17-26.

1905b. On Fishes from the Persian Gulf, the Sea of Oman

and Karachi, Collected by Mr. F. W. Townsend.

Journal of the Bombay Natural History Society,

16:318-333.

1908. Report on the Marine Fishes Collected by Mr. J.

Stanley Gardiner in the Indian Ocean, London.

Transactions of the Linnean Society of London,

12:217-255, plates 23-32.

1915. A Collection of Fishes from Lagos. Annals and

Magazine of Natural History, series 8, 15:124-130.

1920. Fishes from Durban, Natal, Collected by Messrs

H. W. Bell Marley and Romer Robinson. Annals of

Durban Museum, 2(5): 199-222.

1921a. Three New Fishes from South Africa Collected by

Mr. H. W. Bell Marley. Annals of Durban Museum,

3:1-2.

1921b. New Fishes from Deep Water off the Coast of

Natal. Annals and Magazine of Natural History,

(9)7:412-429.

Rendahl, H.

1921. Results of Dr. E. Mjoberg's Swedish Scientific

Expeditions to Australia. 1910-13, XXVIII: Fische.

Kongliga Svenska Vetenskaps-Akadcmi Handlingar,

61:17-24.

Richardson, J.

1846. Report on the Ichthyology of the Seas of China

and Japan. Report of the Fifteenth Meeting of the

British Association for the Advancement of Science,

1845:187-320.

Rochebrune, A. T. de

1882. Faune de la Senegambie: Poissons. Actes de la

Sociite" linne'enne de Bordeaux, 6:37-180, 6 plates.

Roxas, H. A., and C. Martin

1937. A Check List of Philippine Fishes. Technical Bulletin, Department of Agriculture and Commerce,

Philippine Islands, 6:1-314.

•Ruppell, W. P. E. S.

1826. Atlas Zu der Reise im nordlichen Afrika, Zoologie:

Fische des rothen Meeres. 4 volumes.

Russell, P.

1803. Descriptions and Figures of 200 Fishes, Collected at

Vizagapatnam on the Coast of Coromandal. 2 volumes, 197 plates. London.

Rutter, C. M.

1897. A Collection of Fishes Obtained in Swatow, China,

by Miss Adele M. Fielde. Proceedings of the

Academy of Natural Sciences of Philadelphia, 1897:

56-90.

Said, R.

1962. The Geology of Egypt. 377 pages. Amsterdam &

New York: Elsevier Publishing Co.

Saramma, A. D.

1963. Bottom Fishes Collected by the Research Vessel

Conch, off the Kerala Coast during 1958-63:

Heterosomata. Bulletin of the Department of Marine

Biology and Oceanography (University of Kerala),

1:57-80.

Sauvage, H. E.

1878. Cynoglossus (Arelia) solum sp. nov. from Me-Kong.

Bulletin de la Sociiti Philomathique de Paris, 7:92-

95.

1881. Sur une collection de Poissons de Swatow (S. China).

Bulletin de la Sociiti Philomathique de Paris,

(7)5:104-107.

1883. Sur une collection de poissons recudllie dans le
NUMBER 2S8 107

Me-nam (Siam) par M. Harmand. Bulletin de la

Soditi Philomathique de Paris, (7)7:150-155.

Schmidt, P., and G. Lindberg

1930. A List of Fishes, Collected in Tsuruga (Japan) by

Dr. W. Roszkowski. Bulletin V Academy Science

I'URSS, 1930:1135-1150.

Schneider, J. G.

1801. Systema Ichthyologiae. ix + 584 pages, 110 plates.

Schultze, O.

1897. Weber herstellung und conservirung durchaichtige

ombryonen zum studium der skeletbildung. Anatomischer Anzeiger, 13:3-5.

Scott. J. S.

1959. An Introduction to the Sea Fishes of Malaya.

XII + 180 pages. Kuala Lumpur.

Scale, A. V.

1910. Fishes of Borneo with Description of Four New

Species. Philippine Journal of Science, 5:263-289,

4 plates.

1914. Fishes of Hong Kong. Philippine Journal of Science,

9:59-81.

Seshappa, G., and B. S. Bhimachar

1955. Studies on the Fisheries and Biology of the Malabar

Sole, Cynoglossus semifasdatus Day. Indian Journal

of Fisheries, 2(1): 180-230, 1 plate, 21 figures.

Sewell, R. B. S.

1948. The Free-swimming Planktonic Copepods, Geographical Distribution. Pages 317-502 in volume 8

of Scientific Reports, John Murray Expedition. 317—

502.

Shen, S.

1967. Studies on Fiat-Fishes (Pleuronectiformes or Heterosomata) in the Adjacent Waters of Hongkong.

Quarterly Journal of the Taiwan Museum, 20(1,2):

150-281, 160 figures.

1969. Additions to the Study on the Fiat-Fishes in the

Adjacent Waters of Hongkong. Report, Institute of

Fisheries Biology (Taiwan), 2:19-27.

Smith, H. M.

1933. Contributions to the Ichthyology of Siam, IV:

Fishes Not Previously Recorded from Siam. Journal

of the Natural History Society of Siam, supplement,

9(l):53-87, 3 plates, 4 figures.

1945. The Fresh Water Fishes of Siam, or Thailand.

Bulletin of the United States National Museum,

188:I-XII + 1-622, 9 plates, 107 figures.

Smith, H. M., and T. C. B. Pope

1906. List of Fishes Collected in Japan in 1903, with

Descriptions of New Genera and Species. Proceedings

of the United States National Museum, 31:459-499,

12 figures.

Smith, J. L. B.

1949[1965]. The Sea-Fishes of Southern Africa. 580 pages,

107 plates, 1219 figures. Cape Town: Cape and

Transvaal Printers Ltd.

Snyder, J. O.

1909. Description of New Genera and Species of Fishes

from Japan and the Riu Kiu Islands. Proceedings

of the United States National Museum, 36:597-610.

1912. Japanese Shore Fishes Collected by the United States

Bureau of Fisheries Stamer "Albatross" Expedition

of 1906. Proceedings of the United Slates National

Museum, 42:399-450, 11 plates.

Sowerby, A. de C.

1930. Ihe Freshwater and Estuarine Fishes of Manchuria:

The Fishes of the Manchurian Seas. Naturalist in

Manchuria, 4:43-286, figures.

Stauch, A. F.

1966. Description d'uno nouvelle espice de Cynoglossidae.

Bulletin du Museum National d' Histoire Naturelle

de Paris, (2)38:126-128.

Steindachner, F.

1867. Uber einige neue und Seltene Meeresfische aus

China. Sitzungsberichte der Akademie der Wissenchaften, Wien, 55:585-592.

1870. Zur Fischfauna des Senegal. Sitzungsberichte der

Akademie der Wissenchaften in Wien, 60(l):945-995.

1882. Beitrage zur Kenntniss der Fische Afrika't und

Beschreibung einer newen Paraphoxinus—Art aus

der Herzegowina. Denkschriften der Akademie der

Wissenchaften, Wien, 45(1): 1-18, 6 plates.

1894. Die Fische Liberia's. Notes from the Ley den Museum, 16:1-96.

1896. Bericht uber die Wahrend der Reise Sr. Maj. Schiff

"Aurora" von Dr. C. Ritter V. Microszewski in den

Jahren 1895 und 1896, gesammelten Fische. Annalen

des Naturhistorischen Museums in Wien, 11:197-230.

•1898. Reise Szechenyi Ostasien. Volume 2.

Steinitz, W.

1927. Beitrage zur Kenntnis der Kustenfauna Palastinas.

Erster Teil in Pubblicazioni della Stazione Zoologica

di Napoli, 8(3-4):311-353.

1929. Die Wanderung indopazifischer artenins Mittelmeer

seit Beginn der Quartarperiode. International Review of Hydrobiology, 22(12): 1-90.

Suvatti, C.

1936. Index to Fishes of Siam. 226 pages, 1 plate, 1 chart.

Bangkok: Bureau of Fisheries.

1950. Fauna of Thailand. 1100 pages. Bangkok: Department of Fisheries, Thailand.

Taylor, W. R.

1967. An Enzyme Method of Clearing and Staining Small

Vertebrates. Proceedings of the United Stales National Museum, 122:1-17.

•Tchang, T. L.

1929. A Review of the Fishes of Nanking. Contributions

from Biological Laboratory of Science Society of

China, Zoological Series, volume 4, number 4.

1955. Fishes of the Yellow Sea and Po-hai, China. 353

pages.

•Thompson, W. W.

1918. Report of the Marine Biologist. Volume 4. Cape

Town.

Tortonese, E.

1964. The Main Biogeographical Features and Problems

of the Mediterranean Fish Fauna. Copeia, 1964(1):

98-107.

Ui, N.

1929. Monograph of Fishes from Province Kii. 284 pages.

Vaillant, L. L.
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1893. Contribution a letude de la faune ichthyologique de

Borneo. Nouvelles Archives du Museum National

d'Histoire Naturellc de Paris, 3(5):23-112. 2 plates.

Vinciguerra, D.

1889. Viaggio-di Leonardo Fea in Birmania e regioni

Vicine (Pesci). Annali del Museo Civico di Storia

Naturale di Genova, 2(9): 129-362.

Vortsman, A.

1927. Otolithus (Solea) botanggensis and Otolithus (Solea)

sonogonneisis from Eocene of Java. Wettenschappen

Mededeenlingen, 5:13-14. figures 16-17.

Waite, E. R.

1905. Notes on Fishes from Western Australia. Record

of the Australian Museum, 6:55-82.

Wang, K. F.

1933. Study of the Teleost Fishes of Coastal Region of

Shangtung. Part I in Contributions from the Biological Laboratory of the Science Society of China,

Nanking, Zoological Series, 9(1): 1-76, 39 figures.

Weber, M.

1910. Nuee Fische aus Niederlandisch Sud-Neu-Guinea.

Notes from the Leyden Museum, 32:225-240.

1913a. Susswassernsche aus Niederlandische Sud-und NordNew Guinea. Pages 513-613 in part 4 of volume 4

in Zoologie in Nova Guinea. Resultats de V Expedition Scientifique Neerlandaise a la Nouvella-Guinee.

3 plates.

1913b. Die Fische der Siboga-Expedition. Siboga Expedition Report, 57:1-710.

Weber, M., and L. F. de Beaufort

1929. The Fishes of the Indo-Australian Archipelago.

Volume 5, XIV + 458 pages, 98 figures. Leiden:

E. J. Brill Ltd.

Whitehead, P. J. P.

1969. The Reeves Collection of Chinese Fish Drawings.

Bulletin of the British Museum (Natural History),

Zoology, 3(7): 13-233. 20 plates.

Woodward, A. S.

1910. On a Fossil Sole and Sole Eel from the Eocene of

Egypt. Geological Magazine, 7:402-405. 1 plate.

Wu. H. W.

1929. Study of Fishes of Amoy. Contributions from the

Biological Laboratory of the Science Society of

China, 5(4):l-90.

1931. Note sur les poissons mar ins recueillis par M. Y.

Chen sur la cdte du Tch&iang avec synopsis des

especes du genre Tridentiger. Sinensia Nanking,

1:165-174.

1932. Contributions a' I'itude morphologicue, biologique

et systematique des poissons heterosomes (Pisces,

Heterosomata) de le Chine. 178 pages, 27 figures.

Thesis, University of Paris.

1933. Study of the Teleost Fishes of Coastal Region of

Shungtung. Contributions of Biological Laboratory

of Science Society of China, Zoological Series. 9(1):

37-65.

Wu, H. W., and K. F. Wang

1933. Supplementary Notes on the Fishes of Heterosomata

of China. Contributions from the Biological Laboratory of the Science Society of China, Zoological

Series, 9(7):297-304, 1 figure.

Yanai, T.

195O.The Fishes of the Sanin District. Zoological Magazine

(Tokyo), 59:17-22.

Zeuner, F. E.

1959. The Pleistocene Period: Its Climate, Chronology, and

Faunal Succession, 447 pages. London: Hutchinson

& Co. Ltd.
NUMBER 238 109

PLATE 1.—Cynoglossus canariensis Steindachner: a, lateral view of ocular side of specimen 325

mm SL (USNM 188435) from Lagos coast, Nigeria; b, enlarged view of anterior part of same.

Cynoglossus monodi Chabanaud: c, lateral view of ocular side of specimen 314.0 mm SL (USNM

188434) from Lagos coast, Nigeria; d, enlarged view of anterior part of same.
110

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 2.—Cynoglossus senegalensis (Kaup): a, lateral view of ocular side of specimen 355.0 mm

SL (USNM 247424) from Lagos coast, Nigeria; b, enlarged view of anterior part of same.

Cynoglossus dubius Day: c, lateral view of ocular side of specimen 370.0 mm SL (SOSC A 23)

from Arabian Sea; d, enlarged view of anterior part of same.
NUMBER 238 111

PLATE 3.—Cynoglossus borneensis (Bleeker): a, lateral view of ocular side of specimen 271.0

mm SL (CAS., Vanderbilt collections, Reg 2582) from Shakon Province, yt miles off shore,

Gulf of Thailand; b, enlarged view of anterior part of same. Cynoglossus browni Chabanaud:

c, lateral view of ocular side of specimen 305.0 mm SL (USNM 188437) caught off the coast of

Nigeria; d, enlarged view of anterior part of same.
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 4.—Cynoglossus bilineatus (Lacepede): a, lateral view of ocular side of specimen 310.0

mm SL from Hong Kong; b, enlarged view of anterior part of same. Cynoglossus attenuatus

Gilchrist: c, lateral view of ocular side of specimen 244.0 mm SL (ANSP 88867) from Delagoa

Bay, South Africa; d, enlarged view of anterior part of same.
NUMBER 238 113

PLATE 5.—Cynoglossus lachneri Menon: a, lateral view of ocular side of specimen 226.0 mm SL

from Silhoutte Bay, Seychelles; b, enlarged view of anterior part of same. Cynoglossus dispar

Day; c, lateral view of ocular side of specimen 303.0 mm SL (SOSC A 23) from Karachi, west

of Astola Island, Pakistan, lat. 23°06'35"N, long. 63°48'65"E; d, enlarged view of anterior

part of same.
114

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 6.—Cynoglossus kopsi (Bleeker): a, lateral view of ocular side of specimen 178 mm SL

(USNM 137982) from China Sea, near Hong Kong; b, enlarged view of anterior part of same.

Cynoglossus joyneri Giinther: c, lateral view of ocular side of specimen 166.0 mm SL (KU

26965) from Omuta, Fukuoka Prefecture, Japan; d, enlarged view of ocular side of anterior part

of same.
NUMBER 238

115

m

PLATE 7.—Cynoglossus itinus (Snyder): a, lateral view of ocular side of specimen 124.0 mm SL

(USNM 72023) from Tanegashima, Japan; b, enlarged view of anterior part of same. Cynoglossus maculipinnis Rendahl: c, lateral view of ocular side of specimen 102.0 mm SL (WAM P

11729) of C. maculipinnis from Exmouth Gulf; d, enlarged view of anterior part of same.
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

PLATE 8.—Cynoglossus broadhursti Waite: a, lateral view of ocular side of specimen 214.0 mm

SL (WAM P 14741) from Shark Bay, Western Australia; b, enlarged view of anterior part of

same. Cynoglossus ecaudatus Gilchrist: c, lateral view of ocular side of specimen 150.0 mm SL

(SOSC Ref. No. 145) from Somali coast; d, enlarged view of anterior part of same.
NUMBER 238 117

PLATE 9.—Cynoglossus cadenati Chabanaud: a, lateral view of ocular side of holotype 145.0 mm

SL (MNHP 49.20) from Senegal, west Africa; b, enlarged view of anterior part of same. Cynoglossus sinusarabici Chabanaud: c, lateral view of specimen 83.0 mm SY (MNHP 1967.600);

d, enlarged view of ocular side of anterior part of same.
118

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 10.—Cynoglossus sealarki Regan: a, lateral view of ocular side of paralectotype 182.0

mm SL (BMNH 1908.23.155) from Saya-de-malha Bank; b, enlarged view of anterior part of

same. Cynoglossus zanzibarensis Norman: c, lateral view of ocular side of specimen, 154.0 mm

SL (SOSC Ref. No. 170) from Mozambique coast, lat. 25°12'S, long. 4O°S1'£; d, enlarged view

of anterior part of same.
NUMBER 238 119

PLATE 11.—Cynoglossus capemis (Kaup): a, lateral view of ocular side of neotype 167.0 mm SL

(BMNH 1897.10.8.2) from Saldanha Bay; b, lateral view of anterior part of same. Cynoglossus

arel (Schneider): c, lateral view of ocular side of specimen 348.00 mm SL (SOSC Ref 334)

from Porto Novo, Madras, lat. 11°32'N, long. 79853'E; d, enlarged view of ocular side of anterior part of same.
120

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 12.—Cynoglossus robustus Gunther: a, lateral view of ocular side of specimen 340.0 mm

SL (USNM 56386) from Japan; b, enlarged view of ocular side of anterior part of same.

Cynoglossus lingua (Hamilton-Buchanan): c, lateral view of ocular side of specimen 315.0 mm

SL (CAS George Vanderbilt collection 2582) from Gulf of Thailand; d, enlarged view of anterior part of same.
NUMBER 238 121

PLATE 13.—Cynoglossus cynoglossus (Hamilton-Buchanan); a, lateral view of ocular side of specimen 116.0 mm SL (ZSI 1499) from Calcutta; b, enlarged view of anterior part of same. Cynoglossus semifasciatus Day: c, lateral view of ocular side of specimen 114.0 mm SL (SOSC Ref

334) caught off Thirumulli Vasal Village, near Porto Novo, Madras; d, enlarged view of anterior

part of same.
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 14.—Cynoglossus macrostomus Norman: a, lateral view of ocular side of specimen 127.0

mm SL (SOSC Ref 334) from Neendakari, Kerala State; b, enlarged view of anterior part of

same. Cynoglossus tnonopus (Bleeker): c, lateral view of ocular side of specimen 147.0 mm SL

(GVF collection) from Gulf of Thailand; d, enlarged view of anterior part of same.
NUMBER 238 123

- • •

PLATE 15.—Cynoglossus puncticeps (Richardson): a, lateral view of ocular side of specimen 105.0

mm SL (GVF) from Gulf of Thailand; b, enlarged view of anterior part of same. Cynoglossus

durbanensis Regan: c, lateral view of ocular side of specimen 153.0 mm SL (ANSP 63890) from

Durban; d, enlarged view of anterior part of same.
124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 16.—Cynoglossus gilchristi Regan: a, lateral view of ocular side of specimen 90.5 mm SL

(SOSC Ref 134) from Nossi Be, Madagascar; b, enlarged view of anterior part of same.

Cynoglossus lida (Bleeker): c, lateral view of ocular side of specimen 180.0 mm SL (SOSC Ref

170) from Mozambique coast, lat. 19°09'S, long. 36°20'E; d, enlarged view of anterior part of

same.
NUMBER 238

PLATE 17.—Cynoglossus carpenteri Alcock: a, lateral view of ocular side of specimen 179.0 mm

SL (SOSC A 23) from Arabian sea, lat. 24°13'N, long. 65°52'E; b, enlarged view of anterior part

of same. Cynoglossus acutirostris Norman: c, lateral view of ocular side of paralectotype 190.0

mm TL (USNM 109493) from the Gulf of Aden; d, enlarged view of anterior part of same.
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 18.—Cynoglossus marleyi Regan: a, lateral view of ocular side of specimen 159.0 mm SL

(SOSC) caught off Natal. Cynoglossus suyeni Fowler: c, lateral view of ocular side of specimen

247.0 mm SL (USNM 137944) from the Philippine-Celebes seas; d, enlarged view of anterior

part of same.
NUMBER 238 127

d

PLATE 19.—Cynoglossus heterolepis Weber: a, lateral view of ocular side of specimen 138.0 mm

SL (USNM 174028) from Northern Territory, Australia; b, lateral view of anterior part of same.

Cynoglossus waandersi (Bleeker): c, lateral view of ocular side of specimen 73.0 mm SL (UMMZ

171680) from Morsi River at Mooara, Sumatra; d, lateral view of anterior part of same.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 20.—Cynoglossus semilaevis Giinther: a, lateral view of ocular side of specimen 350 mm

SL (BMNH 1939.1.17.56) from Hong Kong; b, enlarged view of anterior part of same. Cynoglossus abbreviatus Giinther: c, lateral view of ocular side of specimen 270.0 mm SL (USNM)

from Pearl River mouth, Hong Kong; d, enlarged view of anterior part of same.
NUMBER 238 129

PLATE 21.—Cynoglossus gracilis Giinther: a, lateral view of ocular side of specimen 180.0 mm SL

(USNM 86466) from China. Cynoglossus microlepis (Bleeker): b, lateral view of ocular side of

specimen 215.0 mm SL (UMMZ 181238) from Cambodia; c, enlarged view of anterior part of

same. Cynoglossus macrophthalmus Norman: d, lateral view of anterior half of ocular side of

specimen 70.0 mm SL (AM 1A 6460) from Lindeman Island, Queensland, Australia.



REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION

Manuscripts intended for series publication receive substantive review within their

originating Smithsonian museums or offices and are submitted to the Smithsonian Institution

Press with approval of the appropriate museum authority on Form SI—36. Requests for

special treatment—use of color, foldouts, casebound covers, etc.—require, on the same

form, the added approval of designated committees or museum directors.

Review of manuscripts and art by the Press for requirements of series format and style,

completeness and clarity of copy, and arrangement of all material, as outlined below, will

govern, within the judgment of the Press, acceptance or rejection of the manuscripts and art.

Copy must be typewritten, double-spaced, on one side of standard white bond paper,

with l%" margins, submitted as ribbon copy (not carbon or xerox), in loose sheets (not

stapled or bound), and accompanied by original art. Minimum acceptable length is 30 pages.

Front matter (preceding the text) should include: title page with only title and author

and no other information, abstract page with author/title/series/etc., following the established format, table of contents with indents reflecting the heads and structure of the paper.

First page of text should carry the title and author at the top of the page and an unnumbered footnote at the bottom consisting of author's name and professional mailing address.

Center heads of whatever level should be typed with initial caps of major words, with

extra space above and below the head, but with no other preparation (such as all caps or

underline). Run-in paragraph heads should use period/dashes or colons as necessary.

Tabulations within text (lists of data, often in parallel columns) can be typed on the text

page where they occur, but they should not contain rules or formal, numbered table heads.

Formal tables (numbered, with table heads, boxheads, stubs, rules) should be submitted as camera copy, but the author must contact the series section of the Press for editorial attention and preparation assistance before final typing of this matter.

Taxonomic keys in natural history papers should use the aiined-couplet form in the

zoology and paleobiology series and the multi-level indent form in the botany series. If

cross-referencing is required between key and text, do not include page references within the

key, but number the keyed-out taxa with their corresponding heads in the text.

Synonymy in the zoology and paleobiology series must use the short form (taxon,

author, yearpage), with a full reference at the end of the paper under "Literature Cited."

For the botany series, the long form (taxon, author, abbreviated journal or book title, volume,

page, year, with no reference in the "Literature Cited") is optional.

Footnotes, when few in number, whether annotative or bibliographic, should be typed

at the bottom of the text page on which the reference occurs. Extensive notes must appear at

the end of the text in a notes section. If bibliographic footnotes are required, use the short

form (author/brief title/page) with the full reference in the bibliography.

Text-reference system (author/year/page within the text, with the full reference in a

"Literature Cited" at the end of the text) must be used in place of bibliographic footnotes in

all scientific series and is strongly recommended in the history and technology series:

"(Jones, 1910:122)" or ". . . Jones (1910:122)."

Bibliography, depending upon use, is termed "References," "Selected References," or

"Literature Cited." Spell out book, journal, and article tities, using initial caps in all major

words. For capitalization of titles in foreign languages, follow the national practice of each

language. Underline (for italics) book and journal titles. Use the colon-parentheses system

for volume/number/page citations: "10(2):5-9." For alinement and arrangement of

elements, follow the format of the series for which the manuscript is intended.

Legends for illustrations must not be attached to the art nor included within the text but

must be submitted at the end of the manuscript—with as many legends typed, doublespaced, to a page as convenient.

Illustrations must not be included within the manuscript but must be submitted separately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.)

can be intermixed throughout the printed text. They should be termed Figures and should

be numbered consecutively. If several "figures" are treated as components of a single larger

figure, they should be designated by lowercase italic letters (underlined in copy) on the illustration, in the legend, and in text references: "Figure 9b." If illustrations are intended to be

printed separately on coated stock following the text, they should be termed Plates and any

components should be lettered as in figures: "Plate 9b." Keys to any symbols within an

illustration should appear on the art and not in the legend.

A few points of style: (1) Do not use periods after such abbreviations as "mm, ft,

yds, USNM, NNE, AM, BC." (2) Use hyphens in spelled out fractions: "two-thirds." (3)

Spell out numbers "one" through "nine" in expository text, but use numerals in all other

cases if possible. (4) Use the metric system of measurement, where possible, instead of

the English system. (5) Use the decimal system, where possible, in place of fractions.

(6) Use day/month/year sequence for dates: "9 April 1976." (7) For months in tabular listings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc.

Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front

matter and ALL legends, etc., at the back of the text—in the following order: (1) title page,

(2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes,

(7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.
K

1

•

"iHr

^ ^MBI