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NELSON~Gareth Jon~ 193'7-
GILL ARCHES OF TELEOSTEAN FISHES.
University of Hawaii~ Ph.D.~ 1966
Zoology
UniversityMicrofilms, Inc., Ann Arbor, Michigan
GILL ARCHES OF
TELEOSTEAN FISHES
A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE
UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT
OF THE REQ,UIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
IN ZOOLOGY
JUNE 1966
By
Gareth Jon Nelson
Thesis Committee:
Dr. Albert L. Tester, Chairman
Dr. George W. Gillett
Dr. Sidney C. Hsiao
Drc Barry S. Muir
Dr. Ernst S. Reese
iii
ABSTRA.CT
Gill arches were examined for some osteoglossiform,
clupeiform, anguilliform, cypriniform, scopeliform,
atheriniform, percopsiform, gadiform, beryciform, zeiform,
syngnathiform, gasterosteiform, perciform, pleuronectiform,
scorpaeniform, echeneiform, tetraodontiform and
symbranchiform fishes. Arch modifications in generalized
forms are reasonably represented in the clupeiform
scopeliform-beryciform-perciform main line of teleostean
evolution. Arches of elopids are the most primitive and
,
generalized in having suprapharyngobranchials in the first
and second arches and numerous dermal elements not fused
with their endochondral supports. Arches in other groups
may have been derived. from a stage like that of the Recent
Elops by reduction, loss or fusion of certain arch elements.
These modifications form. a trend toward the development of
movable, toothed surfaces, above and below, at the posterior
limit of the pharynx and the loss of toothed surfaces
elsewhere within the bucco-pharyngeal cavity.
Muscles attached to arches were examined in Elops,
Polymixia, Epinephelus. Muscle modifications form a trend
toward the development of longitudinal muscles extending
between the toothed third infrapharyngobranchials and
vertebral column above, and t he toothed fifth cerat
0
branchials and the basibranchial series or urohyal below.
iv
Arch and muscle modifications together constitute a basic
advance in the teleostean feeding mechanism appearing at the
scopeliform level of structural organization.
Arches were examined for representatives of the percoid
families Girellidae, Embiotocidae, Pomacentridae, Labridae
and Scaridae. The series Girella-ChromiS-Embioto~­
Bodianus-Scarus is one of increasing structural
specialization and is consistent with t he hypothesis that
the groups represented are closely related.
Arches were examined for representatives of eighteen
eel families. On the basis of gill arch and other
characters the eels may be divided into three evolutionary
lineages: anguilloid (Anguillidae, Heterenchelidae,
Serrivomeridae, Nemichthyidae, Moringuidae, Xenocongridae,
Dysomminidae, Muraenidae), synaphobranchoid
(Synaphobranchidae, Ilyophidae, Simenchelidae, Dysommidae),
congroid (Congridae, Heterocongridae, Nessorhamphidae,
Nettastomidae, Derichthyidae, Ophichthidae, Muraenesocidae).
In each lineage arches tend to be similarly modified in
relation to body form and food habits. Gills are
posteriorly displaced in the more elongate forms. Loss of
arch elements and muscle modifications result in some cases
in a highly expandible pharynx, probably an adaptation
enabling the fishes to swallow large prey. ftPharyngeal jawsft
of muraenids seem to have developed in relation to mechanical
problems of transporting prey through a secondarily
v
elongate pharynx.
Eels of the genus Neenchelys are referred to the family
Ophichthidae, subf~ily Echelinae, on the basis of numerous
and overlapping branchiostegal rays, structure of the
sensory canal system of the head, cranial osteology and a
well developed caudal fin.
The family Muraenidae is divid~d into two sUbfamilies,
Uropterygiinae (new sUbfamily) and Muraeninae.
Arches were examined for representatives of the
families Chirocentridae, Engraulidae, Clupeidae and Chanidae.
Arch modifications in clupeids include the loss of teeth,
proliferation of gill rakers and development of an
epibranchial organ.
Osteoglossids, clupeids, Chanos, Curimata,
Alepocephalus and other forms were examined for epibranchial
organs. These organs seem always to be formed by a
secondary upgrowth of the fifth gill slit posterior to the
fifth epibranchial into a developing blind diverticulum of
the pharynx. They apparently have independently developed
in at least four evolutionary lineages: osteoglossid,
characid, clupeid and chanid.
TABLE OF CONTENI'S
Abstract iii
• • • • • • • • • • • • • • • • • • • • • • •
..
List of tables ix
• • • • • • • • • • • • • • • • • • •
List of figures • • • • • • • • • • • • • • • • • • :x
0
SECTION I INTRODUCTION • • • • • • .. • • • • • • • • 1
.
1.1 Scope of the present work .. • • • • • • • .. .. 1
1.2 Scope of previous work on the visceral
apparatus of fishes • • • • • • • • • • • • • J
Sources of material and acknowledgments • • • • 7
SECTION II MATERIAL AND METHODS • .. • .. • •• • • • • 10
SECTION III MAJOR FEATURES OF THE VISCERAL APPARATUS
OF A GENERALIZED TELEOSTEAN FISH • • 20
• •
SECTION IV THE BRANCHIAL APPARATUS IN GENERALIZED
TELEOSTEAN FISHES .. 22
• • • • • • • • • • •
4.1 Explanation of terminology, figures and format 22
4.1.1 Terminology .. 22
• • • • • • • • • • • • • •
4.1.2 Figures • • • • • • • • • • • • • • • • • 24
4.1.3 Format • • • • • • • • • • • .. • • • • • 25
. .
·
4.2 Gill arches. .. • • 0 ;; • • • • • • • • • 39
4.2.1 Elopoid fishes • • • • • • • • e • • • • 39
4.2.2 Osteoglossiform fishes • • • • • • • • • 48
4.2.3 Cypriniform fishes 52
• • • • • • • • • • •
Salmonoid fishes 54
• • • • 0 • • • II • • •
· . . . . . . . . . . .
Esocoid fishes 56
;
Scopeliform fishes • • • • • • • • • • • 57
vii
4.2.7 Beryciform fishes 58
• • • • • • • • • • • •
Perciform. fishes • • 59 .
• • • • • • • • • 0 •
Discussion • • • 64
• • • • • • • • • • • • •
4.3 Arteries associated with gill arches • • 72
• • • •
Introductory remarks • • • • • • • 72
• fit • •
4.3.2 Arteries of Elops hawaiiensis • • • • • • 73
4.3.3 Concluding remarks • • • • • • • • • • • • 78
4.4 Muscles attached to the gill arches .. 80
• • • • •
4.4.1 Elops, Polymixia, Epinephelus • • • • • • 80
4.4.2 Discussion 83
• • • • • • • • • • • • • • • •
4.5 Functional significance of gill arch and
muscle modifications 89
• • • • • • • • • • • • •
SECTION V THE BRANCHIAL APPARATUS OF TELEOSTEAN FISHES
OF THE ORDER ANGUILLIFORMES .. • • • • • • • 92
Introduction • • 92
• • • • • • • •
• s • • • • • •
5.2 Gill arches 92
• • • • • • • • • • • • • • • • • •
5.2.1 Results 92
• • • • • • • • • • • • • • • • •
Discussion • • • • • • • • • • • • • • • • 100
5.3 Muscles attached to the gill arches 116
• • • • • •
5.3.1 Results 116
• • • • • • • • • • • • • • • • •
Discussion • 122
• • • • • • • • • • • • • • •
5.4 Functional significance of gill arch
modifications 124
• • • • • • • • • • • • • • • • •
5.5 Remarks on eel origins • • 131
• • • • • • • • • • •
5.6 Addendum: Osteology and relationships of the
eel Neenchelys buitendi1!1 • • • • • • • • •• 133
viii
5.6.1 Introduction ••••••••••••••• 133
5.6.2 Osteology •••••••••••••••• 134
5.6.3 Discussion •••••••• 136
0 •••••••
SECTION VI THE BRANCHIAL APPARATUS OF TELEOSTEAN FISHES
OF THE SUBORDER CLUPEOIDEI • • • • • • • • 142
6.1 Preliminary remarks ••••• 142
0 ••••••••
6.2 Gill arches •••••••••••••••••• 143
6.2.1 ResuIts ••••••••••••• • -. • • 143
6.2.2 Discussion ••••••••••••••• 149
0
6.3 Epibranchial organs •••••••••••••• 158
6.3.1 Introduction ••••••••••••••• 158
6.3.2 Results ••••••••••••••••• 160
6.3.3 Discussion •••••••••••••••• 165
6.4 Concluding remarks ••••••••••••••• 168
SECTION VII SUMMARY • • • • • • • • • • • • • • • • • 172
7.1 Clupeiform-scopeliform-beryciform-perciform
series ••••••••••••••••••• • 172
7.G Anguilliformes • • • • • • • • • • • • • • • • • 175
7.3 Clupeoidei ••• 177
0 •••••••••••••••
Bibliography • 180
0 • • • • • • • • • • • • • • • • • • •
Figures 1-250 • • • • • • • • • • • • • 200
0 • • • • • • •
ix
LIt."'T OF TABLES
Table I. Summary of enCl.Jchondral gill arch elements
in generalized repr~sentatives of some
teleostean groups - - - - - - - - - - - - - 26
Table II. Summary of some dermal elements in some
teleostean fishes - - - - - - - - - - - - - - 27
Table III. Summary of endochondral gill arch elements
in osteoglossiform fishes - - - - - - - - - - - - - 51
Table IV. Summary of selme endochondral and dermal
gill arch elements in representatives of the
perciform families Girellidae, Pomacentridae,
Embiotocidae, Labridae and Scaridae - - - - - - 62
Table V. Summary of ventral gill arch muscles in
- - - - -
some generalized t~leostean fishes - 81
Table VI. Summary of dorsal gill arch muscles in
some generalized teleostean fishes - - - - - - 82
Table VII. Summary of some gill arch elements in 18
eel families ------- - - - - - - - - 94
Table VIII. Muscles attached to the ventral parts of
the gill arches in some eels - - - - - - - - - 117
Table IX. Muscles attached to the dorsal parts of
the gill arches in some eels - - - - - - - - - 118
Table X. Muscle terminology for ventral gill arch
muscles in Anguilla - - - - - - - - - - - - - - - - 120
Table XI. Muscle terminology for dorsal gill arch
muscles in Anguilla - - - - - - - - - - - - - - 121
Table XII. Measurement data concerning body
elongation and branchial apparatus position in
some eels - - - - - - - - - - - - - - - - - - - - - 127
Table XIII. Measurement data from the literature
of some eel body proportions - - - - - - - - - 141
Table XIV. Summary of some dermal gill arch elements
in clupeoid fishes - - - - - - - - - - - - - - - - 144
x
LIST OF FIGURES
Figure 1. Diagrammatic cross section through the
pharyngeal region of a primitive bony fish - 200
Figure 2. Elops hawaiiensis, head, lateral view of
left side, showing the visceral apparatus in a
contracted condition - - - - - - - - - - - - - - 201
Figure 3. Elops hawaiiensis, head, lateral view of
left side, showing the visceral apparatus in an
expanded condition - - - - - - - - - - - - - - - 201
Figure 4. Elops hawaiiensis, head, lateral view of
left side, with the jaws and cheek removed,
showing the visceral apparatus in a contracted
condition - - - - - - - - - - - - - - - - - - - 202
Figure 5. Elops hawaiiensis~ head, lateral view of
left side, With the jaws and cheek removed,
showing the visceral apparatus in an expanded
condition - - - - - - - - - - - - - - - - - 202
Figure 6. Elops hawaiiensis, head, anterolateral
view of left side, with the jaws and cheek
removed, showing the visceral app&ratus in an
expanded condition - - - - - - - - - - - - - - - 203
Figure 7. Elops hawaiiensis, head, ventral view,
showing the visceral a.pparatus in a contracted
condition - - - - - - - - - - - - - - - - - 203
Figure 8. Elops hawaiiensis, head, ventral view,
showing the visceral apparatus in an expanded
condition - - - - - .. - - - - - - - - - - - 204
Figure 9. Elops hawaiiensis, head, posteroventral
view, showing the visceral apparatus in an
expanded condition - - - - - - - - - - - - - 204
Figure 10. Elops hawaiiensis, head, lateral view of
left side, with the jaws and cheek removed, the
gill arches separated at the junction between
cerato- and epibranchial and the esophagus
opened, showing the location of the end.Qchondral
elements of the gill arch skeleton - - - - - - - 205
Figure 11. Elope hawaiiensis, basibra.nchial series,
lateral view of left side - - - - - - - - - - - 206