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Bony Fish Skeleton

EJ Hilton, Virginia Institute of Marine Science, Gloucester Point, VA, USA


ª 2011 Elsevier Inc. All rights reserved.

A Hierarchical Approach to the Study of the Skeleton Appendicular Skeleton


Cranial Axial Skeleton Conclusion
Postcranial Axial Skeleton Further Reading

Glossary Durophagy Eating hard-shelled organisms such as


Chondral bone Bone that ossifies in association with coral or mollusks.
a cartilage, through either ossification on the Endoskeleton The skeletal system formed by all skeletal
surface of a cartilaginous precursor (perichondral elements that are nondermal in developmental origin.
bone), as an ossification of the cartilaginous Exoskeleton External support structure of an organism
precursor (endochondral bone), or a combination of formed by all skeletal elements developed from
the two. connection to the dermis, including all dermal bones,
Compound bone A bone that is formed by two or more teeth, scales, and fin rays; mineralized exoskeletons
histologically distinct tissues, often the result of fusion were common and extensive in extinct fishes.
during ontogeny. Membrane bone Bone that ossifies directly within
Dermal bone Bone that ossifies in contact with the ligaments or other tissue not in contact with the dermis
dermis. and separate from cartilage.

The skeleton of bony fishes presents a seemingly endless well as different ecological guilds (Amia and Hiodon as
diversity of forms that reflect the varied life history stra- temperate freshwater fishes and Trachurus as a tempe­
tegies of these organisms. The skeleton provides the rate/tropical marine form), it must be kept in mind that
architectural support for musculature, while at the same these are only three out of more than 30 000 species and
time providing protection for internal organs and sensory do not capture the range of variation found in the diver­
systems. The skeleton of bony fishes is composed of sity of bony fishes.
hundreds of individual bony and cartilaginous elements
that can vary in form, reflective of the function of each
element in an organism. A solid understanding of the
A Hierarchical Approach to the Study of
skeleton and its diversity, therefore, is critical for studies
the Skeleton
of functional morphology and ecomorphology. In addi-
tion, because of the complex nature of the skeleton of Because of the complex nature of the skeleton of bony
bony fishes, it is rich in phylogenetic data and has been fishes, there are many ways in which one can approach
fundamental in the formulation of many compelling the study of the skeleton. For instance, one can study the
hypotheses regarding the evolutionary biology of fishes. exoskeleton as distinct from the endoskeleton. Both the
This article provides a brief overview of the skeletal endoskeleton and exoskeleton can be further broken
structure of actinopterygians and fish-like sarcoptery­ down into various parts, based on variation in the details
gians. By working from illustrations of three taxa of their development, for example. The approach adopted
(bowfin, Amia calva; goldeye, Hiodon alosoides; and here is hierarchical and combines dividing the skeleton
Japanese jack mackerel, Trachurus japonicus), some of the into topographic regions and functional units. The pri­
osteological diversity of bony fishes generally is high­ mary division of the skeleton is the axial and appendicular
lighted. These three taxa represent three phylogenetic regions. Both of these are further divided into regional
levels (i.e., Amia is a nonteleostean actinopterygian, units (e.g., cranial vs. postcranial portions of the axial
Hiodon is a basal teleost, and Trachurus, an acanthopter­ skeleton), which themselves are divided into more refined
ygian, is a derived teleost). Although these three taxa functional units. Other terminology exists for portions of
serve as exemplars for various phylogenetic levels, as the skull, including the chondrocranium or neurocranium

434
The Skeleton | Bony Fish Skeleton 435

(roughly equivalent to braincase as used here) and the the anterior portions of the palatoquadrate). In lungfishes,
dermatocranium (all dermal bones of the skull, including the palatoquadrate (forming the upper jaw) is fused to the
the skull roof, infraorbital bones, jaws, and the opercular ethmoid region, forming the so-called autostylic condition
series). However, the dermatocranium comprises ele­ of jaw suspension that is reflective of durophagy (other
ments from many different regions of the skull and is taxa, including holocephalans and tetrapods indepen­
therefore somewhat unwieldy to discuss in the context dently evolved autostyly). The nasal capsule is typically
of functional units. Another complicating factor is that the a large concavity in the lateral surface of the ethmoid
terminology for individual skeletal elements has changed region that supports the nasal lamellae. The largest bone
dramatically over time and there are many synonyms for of the ethmoid region is typically the mesethmoid, which
several elements that are in current use. forms in the cartilage separating the left and right nasal
capsules. In basal actinopterygians and sarcopterygians,
there are sometimes dermal bones that lie anterior or
Cranial Axial Skeleton dorsal to the mesethmoid. For instance, the rostral bone
forms in the anterior midline of the snout in taxa such as
Four major functional units form the skull of bony fishes Polypterus (bichirs), Amia, and the basal teleost genus Elops
(Figure 1): the braincase, skull roof and infraorbital series, (ladyfishes), and this bone supports the anteriormost por­
opercular series, and splanchnocranium. Although these tion of the cephalic sensory canal system. In other basal
are recognizable as discrete units of the skull, the distinc­ teleosts, there is another dermal bone, the supraethmoid,
tion between some of them may be obscured, particularly which develops dorsal to the mesethmoid and does not
in derived representatives (e.g., the bones of the braincase contain any part of the sensory canal; these bones may fuse
and skull roof in teleostean fishes). to form a compound element (e.g., Hiodon). Ventral to, and
As the name suggests, the braincase of bony fishes often tightly sutured to, the mesethmoid is the vomer.
(Figure 2) is the box surrounding the brain and sense Anteriorly, the mesethmoid and (sometimes) the vomer
organs of the head. The braincase may be divided into form the articulation surface for the upper jaws. Forming
four regions (anterior to posterior): the ethmoid, orbital, the posterior wall of the nasal capsule and the articulation
otic and occipital regions. Early in ontogeny the braincase surface for the palatoquadrate is the lateral ethmoid bone.
is entirely cartilaginous, but in most bony fishes the brain­ In nonteleostean actinopterygians, there is a second more
case becomes well ossified in the adult (exceptions may anterior pair of ethmoid bones, the preethmoids (e.g., in
include some deep-sea fishes, in which the bony skeleton Amia).
is reduced generally, and highly pedomorphic taxa, such The orbital region (which is associated with the eye) is
as the gobioid Schindleria). In basal sarcopterygians (e.g., sometimes also called the sphenoid region, in reference to
coelacanths), there is an intracranial joint between the the sphenoid bone of humans, which lies posterior to the
ethmosphenoid and the otico-occipital units. Virtually, ethmoid region and is associated with the orbit. The domi­
all of the bones of the braincase are chondral, with the nant bone of this region in relatively basal taxa is the
exception of the parasphenoid and vomer, two dermal orbitosphenoid, which ossifies in the cartilage that forms
bones that are tightly associated with the ventral surface the septum between the eyes. This bone is paired in basal
of the braincase, and the intercalar, which is a membrane actinopterygians (e.g., sturgeons, Amia), but may be median
bone in extant taxa. Both the vomer and parasphenoid are in the adult (e.g., basal teleosts) or absent (most acanthop­
primitively toothed bones, although only nonteleostean terygians, which lack a bony or cartilaginous septum).
actinopterygians, osteoglossomorph and elopomorph tel­ The other elements that may be considered part of the
eosts, and a few other more derived teleosts have orbital region include the pterosphenoid, sphenotic, and
parasphenoid dentition (i.e., in most actinopterygians the basisphenoid, all of which are positioned between the
parasphenoid is edentulous). In most basal actinoptery­ orbitosphenoid (if present) and the more posterior otic
gians and some basal teleosts, the vomer is a paired region. As is the case with the orbitosphenoid, these bones
element (e.g., in Hiodon, during early ontogeny the may be lost as independent elements or altogether,
vomer is paired but is fused to form a median element particularly in more derived groups of bony fishes.
in the adult, Figure 2(d)). The parasphenoid is often the The otic region is formed by the bones that are in
largest bone of the braincase, covering much of its ventral contact with the semicircular canals and ampullae of the
surface. The parasphenoid bears lateral projections (the inner ear. This region also forms much of the lateral wall
so-called ascending rami) that often tightly embrace the of the braincase posterior to the orbit. The largest bone of
braincase posterior to the orbit. the otic region typically is the prootic, which forms in
The ethmoid region forms the bones and cartilages that association with the foramen or foramina for portions of
surround the nasal capsules. The ethmoid region also the trigeminal and facial cranial nerves, in some taxa
forms the articulation point for the anterior portions of forming the so-called trigeminofascialis chamber. The
the splanchnocranium (in particular the upper jaws and pterotic lies dorsal to the prootic, and in most teleosts is
436 The Skeleton | Bony Fish Skeleton

dsp pop
(a) io op (b) op
h
le spo
ao dpt le
pto ic
es n
fr pmx epo
n pa pa
fr
r
set soc

es

(c) dpt es (d) dsp


es soc
dsp met
n le fr
n pto exo
ao fr ors
pmx le
io5 pmx ic
io1 pts
io4
smx io4 h
op
mx io3
mpt d enp
sag op
q h
pop
sop mpt
d
ang mx q
pop
rar ar
ang-rar sop
s
pop
op
rar iop
(e) ang
(f) ang-rar

d mx
pmx sop
br
pop
d

Figure 1 Skulls of Amia (a, c, e) and Hiodon (b, d, f) in dorsal (a, b), lateral (c, d), and ventral (e, f) views. Cartilage is shown in dark blue,
chondral bone in purple, dermal bone in light green, compound bone in dark green, and membrane bone in yellow. Abbreviations: ang,
angular; ang-rar, anguloretroarticular; ao, antorbital; ar, articular; br, branchiostegal; dpt, dermopterotic; dsp, dermosphenotic; enp,
endopterygoid; epo, epioccipital; es, extrascapular; fr, frontal; g, gular; h, hyomandibula; ic, intercalar; io, infraorbital; iop, interopercle;
le, lateral ethmoid; met, mesethmoid; mpt, metapterygoid; mx, maxilla; n, nasal; op, opercle; ors, orbitosphenoid; pa, parietal; pmx,
premaxilla; pop, preopercle; pto, pterotic; q, quadrate; r, rostral; rar, retroarticular; s, symplectic; sag, surangular; set, supraethmoid;
smx, supramaxilla; soc, supraoccipital; sop, subopercle. Modified from Grande L and Bemis WE (1998) A comprehensive phylogenetic
study of amiid fishes (Amiidae) based on comparative skeletal anatomy. An empirical search for interconnected patterns of natural
history. Society of Vertebrate Paleontology, Memoir 4: 1–690; and Hilton EJ (2002) Osteology of the extant North American fishes of the
genus Hiodon Lesueur, 1818 (Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142.

incorporated into the surface of the skull roof. In basal fused to form a compound bone, which is often called the
actinopterygians there is an autogenous dermopterotic pterotic. In at least some taxa, these two bones are never
that is a true dermal skull roofing bone that carries a seen as separate elements during ontogeny, and it is often
sensory canal, and is separate from the chondral autop- held as an example of so-called phylogenetic fusion, in
terotic. In teleosts, the autopterotic and dermopterotic are which plesiomorphically separate elements form a single
The Skeleton | Bony Fish Skeleton 437

spo
(a) (b) ors
spo pe dsp dpt
dpt pmx fr ic
le exo
pe dsp ic exo pts pro
fr
pmx pro v boc
le ors pas
pas boc

v pts

soc

pto
epo
es spo
(c) (d)
ic
exo
pto le fr
ic exo
met spo pts pro
fr pts ors
n ors v pas boc
boc
le
bsp pro
v
pas

Figure 2 Bones of the skull roof and braincase of Amia (a, b) and Hiodon (c, d) in lateral (a, c) and ventral (b, d) views. Cartilage is
shown in dark blue, chondral bone in purple, dermal bone in light green, compound bone in dark green, and membrane bone in yellow.
Abbreviations: boc, basioccipital; bsp, basisphenoid; dpt, dermopterotic; dsp, dermosphenotic; epo, epioccipital; es, extrascapular;
exo, exoccipital; fr, frontal; ic, intercalar; le, lateral ethmoid; met, mesethmoid; n, nasal; ors, orbitosphenoid; pas, parasphenoid; pe,
preethmoid; pmx, premaxilla; pro, prootic; pto, pterotic; pts, pterosphenoid; soc, supraoccipital; spo, sphenotic; v, vomer. Modified
from Grande L and Bemis WE (1998) A comprehensive phylogenetic study of amiid fishes (Amiidae) based on comparative skeletal
anatomy. An empirical search for interconnected patterns of natural history. Society of Vertebrate Paleontology, Memoir 4: 1–690 and
Hilton EJ (2002) Osteology of the extant North American fishes of the genus Hiodon Lesueur, 1818 (Teleostei: Osteoglossomorpha:
Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142.

element in derived taxa (this hypothesis however can the posterior semicircular canal of the inner ear.
rarely, if ever, be distinguished from phylogenetic loss). However, it has been shown that at least in extant teleosts,
The bones that interface with the postcranial axial this bone forms in association with cartilage derived from
skeleton and its associated musculature comprise the the occipital arch of early developmental stages.
occipital region of the braincase. The median basioccipi­ Nonteleostean actinopterygians (e.g., sturgeon, gars, and
tal lies in the ventral midline of the occipital region. In Amia) do, however, have an epiotic bone. The intercalar is
basal forms, the basioccipital is posteriorly concave, and found ventral to the epioccipital (in teleosts) or epiotic (in
often resembles a vertebral centrum. This overall similar­ Amia) and in older literature this bone is termed the
ity in form led to the idea that the occipital region of opisthotic. In Amia and extant teleosts, the intercalar is
actinopterygians incorporated the first vertebral centrum composed only of membrane bone. The remaining bone
into its structure. However, there is little structural or of the occipital region in teleostean fishes is the supraoc­
ontogenetic evidence that a centrum is incorporated into cipital (not present in nonteleostean actinopterygians or
the skull in most actinopterygians. A notable exception sarcopterygians). This chondral bone forms in the poster­
is in adult Amia, in which two centra are fused to the ior midline on the dorsal surface of the braincase and may
basioccipital during ontogeny. In some teleosts (e.g., form part of the skull roof. The supraoccipital often bears
Hiodon), in particular the acanthomorphs, the articulation a median crest that extends posteriorly.
between the skull and vertebral column is formed by the The skull roof of bony fishes is formed by those bones
basioccipital and the exoccipitals, a pair of bones that visible in dorsal view of the skull. By this definition bones
form on either side of the foramen magnum. In lungfishes, such as the supraoccipital and the mesethmoid would be
the exoccipital is the only chondral bone of the braincase. considered as part of the skull roof. Plesiomorphically, the
The exoccipital can be readily identified across taxa skull roof comprises only those dermal bones that form the
because it typically contains or defines the foramen for dorsal surface of the skull, and that cover the cartilage and
the vagal nerve. Dorsolateral to the exoccipital are two chondral bones of the braincase. There is a trend across
paired bones, the epioccipital and the intercalar. The the evolution of fishes toward the sinking of the skull
epioccipital is often called the epiotic because it houses roofing bones (i.e., in most basal taxa the bones of the
438 The Skeleton | Bony Fish Skeleton

skull roof are at the surface and are covered by only a thin left and right lower jaws. This gular is medial in most
layer of skin, whereas the bones are deeper and covered by actinopterygians in which it is present, but is paired in
muscle and skin in more derived fishes). In virtually all Polypterus and coelacanths. The interopercle develops in
actinopterygian fishes, these dermal bones include (ante­ the interoperculomandibular ligament and is found only
rior to posterior) the nasals (which may be reduced to in Amia and teleosts among extant fishes. This bone, which
mere tubes surrounding the sensory canal), the frontals, forms a linkage between the opercular series and the
the parietals, and the extrascapulars (as with the nasals, lower jaw, is important for transmission of force during
these may be little more than tubular bones). The frontal jaw opening. The preopercle carries the dorsal cephalic
typically is the largest of the skull roofing bones, followed sensory canal to the lower jaw. In sarcopterygians
by the parietal. In fish-like sarcopterygian fishes, there is (e.g., coelacanths), there are also other bones of the cheek
an additional pair of bones, which has, in part, led to that carry sensory canals (e.g., the squamosal).
controversy regarding the identity of the skull roofing The splanchnocranium includes all the bones and car­
bones between actinopterygians and sarcopterygians. In tilages of the visceral arches, which include the
sarcopterygians, there are paired frontals, parietals, and mandibular, hyoid, and branchial arches (oral jaws,
postparietals. Based on the pattern and association of the hyoid, and gill arches, respectively) as well as the palato­
bones to the sensory canals and their topographic position quadrate (the roof of the mouth). Therefore, although
(relative to the pineal organ), it has been proposed that the they are all part of an integrated unit, they are all treated
bones commonly termed (and which is followed here) the as distinct portions of this region of the skull. The oral
frontals and parietals in actinopterygians are homologous jaws (upper and lower) most frequently bear teeth,
to the parietals and postparietals, respectively, of sarcop­ although entire orders of fishes (e.g., Cypriniformes)
terygians. A change of terminology for actinopterygians may be edentulous. The form of the teeth, as that of the
has been suggested but has not yet been widely adopted. bones of the jaws themselves, reflects the ecology and diet
The infraorbital series of bones (=circumorbital bones) of fishes (e.g., piscivorous fishes often have sharply
is associated with the skull roof by housing the infraorbital pointed teeth set in relatively narrow jaws, whereas dur­
sensory canal, a ventral branch from the supratemporal ophagous fishes have more robust jaws and often have
sensory canal (contained in the pterotic). The infraorbital molariform teeth). The upper jaw (Figures 1 and 3) is
bones define the orbit and are numbered sequentially
(anterior to posterior), although certain elements of the
(a)
series are given particular names (e.g., infraorbital 1,
pmx
which is often enlarged, is called the lachrymal, and the
posteriormost infraorbital, which contains the branching
between the infraorbital canal and the dorsal supraorbital
canal, is called the dermosphenotic). In addition, the ele­
ments that are positioned under the eye are sometimes
termed the suborbitals and those posterior to the eye are
mx
the postorbitals. Basal actinopterygians also have an
antorbital bone, which lies anterior to the orbit. There is (b)
rc
variably a supraorbital bone, which lies dorsal to the orbit,
ap
although this bone does not generally have a connection
to the sensory canal system.
The opercular series forms the bones of the gill-cover smx
and functions in both feeding and respiration. In virtually pmx
all bony fishes, the largest bone of the opercular series is mx
the opercle, which is typically also the dorsal-most of the
series. The only exception is in extant sturgeons and pad­
dlefishes in which the opercle is lost and the large bone
forming the support for the gill cover is the subopercle (the
Figure 3 Bones of the upper jaw of Hiodon (a) and Trachurus
opercle is present in close fossil relatives, but is greatly (b) in lateral view. Cartilage is shown in dark blue and dermal
reduced in size compared to the subopercle). Typically, the bone in light green. Abbreviations: ap, ascending process; mx,
subopercle is smaller and positioned ventral to the opercle, maxilla; pmx, premaxilla; rc, rostral cartilage; smx, supramaxilla.
and bridges the gap between the opercle and the bran­ Modified from Hilton EJ (2002) Osteology of the extant North
chiostegals, which form the supports for the ventral portion American fishes of the genus Hiodon Lesueur, 1818 (Teleostei:
Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology) new
of the opercular flap. In basal actinopterygians, including series 100: 1–142 and Suda Y (1996) Osteology and muscular
the basal teleosts of the family Elopidae (ladyfishes and attachments of the Japanese jack mackerel, Trachurus
tarpons), there is a gular plate, which forms between the japonicus. Bulletin of Marine Science 58: 438–493.
The Skeleton | Bony Fish Skeleton 439

formed by a series of bones: the premaxilla, maxilla, and (a)


supramaxilla (there are sometimes more than one supra­ sag
maxillae positioned along the posterodorsal margin of the ar
maxilla). The premaxilla is the anteriormost of the series par
and articulates with the ethmoid region of the braincase.
In basal forms (e.g., nonteleostean actinopterygians and m c mc rar
basal teleosts), the premaxilla is rigidly articulated with d ang cm
the braincase and is relatively small in comparison to the (b)
maxilla. In these basal forms, the maxilla is the main
tooth-bearing bone of the upper jaw, while the premaxilla
is restricted to the anterior margin of the jaw. Also in cm ar
many early actinopterygians (e.g., Polypterus, palaeonis­ d ang-rar
coids) and sarcopterygians, the maxilla is firmly attached mc
to the other bones of the cheek. In other nonteleostean
actinopterygians and basal teleosts, the maxilla is freed (c)
from the cheek and rotates forward during feeding, form­
ing a mouth that is more round in cross section. Across the
evolution of the teleosts, the premaxilla becomes the
ang-ar
dominant bone of the upper jaw, and the maxilla becomes d
an edentulous bone that is excluded from the margin of
the jaw. With the premaxilla becoming as the primary
mc rar
bone of the upper jaw, exemplified by Acanthomorpha,
also comes the development of a large ascending process Figure 4 Lower jaw of Amia (a), Hiodon (b), and Trachurus (c) in
of the premaxilla. The left and right premaxillae meet in medial view. Cartilage is shown in dark blue, chondral bone in
purple, dermal bone in light green, compound bone in dark green,
the midline of the jaw, and the ascending processes sur­ and membrane bone in yellow. Abbreviations: ang, angular; ang­
round the rostral cartilage, a median cartilage that ar, anguloarticular; ang-rar, anguloretroarticular; ar, articular; c,
articulates with the ethmoid region of the braincase and coronoid; cm, coronomeckelian; d, dentary; m, mentomeckelian;
serves to support the upper jaw during jaw protrusion. mc, Meckel’s cartilage; par, prearticular; rar, retroarticular; sag,
In fact, all of the modifications of the upper jaw seen surangular. Modified from Grande L and Bemis WE (1998) A
comprehensive phylogenetic study of amiid fishes (Amiidae)
across the evolution of teleosts can be correlated to the based on comparative skeletal anatomy. An empirical search for
development of jaw protrusion, which is extremely well interconnected patterns of natural history. Society of Vertebrate
developed in acanthomorph teleosts. Paleontology, Memoir 4: 1–690; Hilton EJ (2002) Osteology of the
The lower jaw of bony fishes is a complex of dermal, extant North American fishes of the genus Hiodon Lesueur, 1818
chondral, compound and membrane-bone elements (Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana
(Zoology) new series 100: 1–142; and Suda Y (1996) Osteology
(Figures 1 and 4). The chondral bones include ossifica­ and muscular attachments of the Japanese jack mackerel,
tions of Meckel’s cartilage, which runs along much of the Trachurus japonicus. Bulletin of Marine Science 58: 438–493.
length of the jaw; in some taxa, it is regressed in adults.
These chondral bones include the mentomeckelian, near
the anterior tip of the jaw, and the articular and retro­ and is typically the largest bone of the lower jaw. In
articular bones at the posterior end of the jaw (there is nonteleostean fishes, there are additional dermal bones,
typically a single articular; Amia is unique in having two which include the surangular on the external surface, the
articular bones). The articular forms the articulation sur­ prearticular on the internal surface (covering much of the
face for the lower jaw with the quadrate, and the posterior region of Meckel’s cartilage), and a series of
retroarticular serves as the attachment site for the inter­ coronoid bones, which are small tooth-bearing elements.
operculomandibular ligament, which contributes to the Therefore, there is a general trend toward loss of bones in
jaw opening mechanism. On the dorsal surface of the lower jaw across actinopterygian fishes.
Meckel’s cartilage is the coronomeckelian, which is a The dorsal portions of the hyoid arch (i.e., the
membrane bone that forms in the tendon of the adductor hyosymplectic cartilage and its ossifications, the
mandibulae muscle where it attaches to the lower jaw. hyomandibila and the symplectic), together with the pala­
Surrounding the chondral portions of the lower jaw is a toquadrate and its ossifications (palatine, ectopterygoid,
series of dermal elements. Two of these, the angular endopterygoid, metapterygoid, and quadrate), form the
(posterior) and the dentary (anterior), are sensory canal- suspensorium or the roof of the mouth (Figure 5). The
bearing bones. In most extant teleosts, the angular is fused palatine, ectopterygoid, and endopterygoid may bear
to either the articular or the retroarticular bone. In most teeth, particularly in relatively basal forms. In nonteleos­
extant fishes, the dentary is the only tooth-bearing bone tean and basal teleostean actinopterygians, there are
440 The Skeleton | Bony Fish Skeleton

(a) articulation of the suspensorium with the ethmoid region


of the braincase. The ectopterygoid and endopterygoid
dpl pal h are dermal bones that form in association with the pala­
op
enp
toquadrate, and form the major bones of the roof of the
mouth. The metapterygoid and quadrate are posterior
mpt ossifications of the palatoquadrate cartilage. The quadrate
sop bears a distinct head for the articulation with the lower
ecp q pop jaw. In teleosts, it also bears a distinct posteroventral
iop
process that is often tightly associated with the symplec­
tic, which is often a stick-like or wedge-like bone that
(b) forms in the ventralmost part of the hyosymplectic carti­
lage. The hyomandibula is the dorsal ossification of the
dpl
hyosymplectic cartilage and is typically the largest bone
of the hyoid arch. This bone forms the posterior articula­
h tion of the suspensorium with the neurocranium as well as
enp
op
the articulation point for the opercle.
The ventral portions of the hyoid arch bridge the
mpt elements of the suspensorium with the branchial arches
ecp (Figure 6). Articulating directly with the suspensorium
q s
pop (typically near the symplectic) is the interhyal, which also
sop articulates with the posterior certatohyal. The anterior
and posterior ceratohyal ossify in a single cartilage, and
iop serve to support the branchiostegals. One (basal taxa) or
two (most teleosts) hypohyal bones lie anteriorly, and
(c)
these articulate with the bones and cartilages of the bran­
chial arches. In Polypterus and teleosts (convergence),
h op there is an urohyal bone, which is a membrane bone that
forms in the tendon of the sternohyoideus muscle. The
urohyal serves to strengthen the attachment of this muscle
pal enp to the branchial arches, which contributes to the rapid jaw
opening that is necessary for suction feeding.
mpt
ecp s pop sop In the midline of the mouth is a series of median ele­
q ments: the basihyal, which is in series with the hyoid arch,
and the basibranchials, which are in series with the bran­
iop chial arches. The basibranchials ossify in an anterior
basibranchial cartilage (copula) and are numbered anterior
Figure 5 Suspensorium and opercular bones of Amia (a), Hiodon to posterior based on their associated branchial arch; a
(b) and Trachurus (c) in lateral view. Cartilage in dark blue, chondral
separate posterior cartilage found in most actinopterygians
bone in purple, and dermal bone in light green. Abbreviations: dpl,
dermopalatine; ecp, ectopterygoid; enp, endopterygoid; h, represents the fourth basibranchial. All of these elements
hyomandibula; iop, interopercle; mpt, metapterygoid; op, opercle; may bear toothplates. The ventral portions of the branchial
pal, palatine; pop, preopercle; q, quadrate; s, symplectic; sop, arches follow the same pattern as the hyoid arch. There is a
subopercle. Modified from Grande L and Bemis WE (1998) A series of hypobranchial elements that articulate directly
comprehensive phylogenetic study of amiid fishes (Amiidae) based
with the basibranchials, and a more distal series of five
on comparative skeletal anatomy. An empirical search for
interconnected patterns of natural history. Society of Vertebrate certaobranchials. There are five ceratobranchials. The
Paleontology, Memoir 4: 1–690; Hilton EJ (2002) Osteology of the fifth ceratobranchial often does not have a dorsal counter­
extant North American fishes of the genus Hiodon Lesueur, 1818 part (i.e., in most fishes there are only four epibranchials)
(Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana and often exhibits extreme modifications in association
(Zoology) new series 100: 1–142; and and Suda Y (1996) Osteology
with the development of the so-called pharyngeal jaws,
and muscular attachments of the Japanese jack mackerel,
Trachurus japonicus. Bulletin of Marine Science 58: 438–493. including hypertrophy and robust molariform dentition
in durophagous fishes (e.g., some members of the family
Sciaenidae). In many fishes, these modifications of the fifth
toothed dermopalatines of varying number. These are ceratobranchial are matched by similar modifications of the
distinct from the palatine (also called the autopalatine), dorsal branchial elements as well.
which is a chondral bone at the anteriormost extent of In some nonteleostean (e.g., sturgeons) and some basal
the palatoquadrate cartilage that forms the anterior teleostean fishes (e.g., Elops), there are three series of
The Skeleton | Bony Fish Skeleton 441

(a) (b)
br
br
ih
chp

cb chp

gr cb
cha gr
cha
hb
hb
cb
hh
bhtp
hh bbtp bb

bb

bbtp

Figure 6 Ventral portion of the hyoid and branchial arches of Amia (a) and Hiodon (b) in dorsal (oral) view. Cartilage is shown in dark
blue, chondral bone in purple, and dermal bone in light green. Abbreviations: bb, basibranchial; bbtp, basibranchial toothplate; bhtp,
basihyal toothplate; br, branchiostegal; cb, ceratobranchial; cha, anterior ceratohyal; chp, posterior ceratohyal; gr, gill raker; hb,
hypobranchial; hh, hypohyal; ih, interhyal. Modified from Grande L and Bemis WE (1998) A comprehensive phylogenetic study of amiid
fishes (Amiidae) based on comparative skeletal anatomy. An empirical search for interconnected patterns of natural history. Society of
Vertebrate Paleontology, Memoir 4: 1–690 and Hilton EJ (2002) Osteology of the extant North American fishes of the genus Hiodon
Lesueur, 1818 (Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142.

elements in the dorsal portions of the branchial arches: ceratobranchials. In these taxa, the supra- and/or infraphar­
the suprapharyngobranchials, infrapharyngobranchials, yngobranchials articulate with the neurocranium. In most
and the epibranchials, which articulate distally with the extant teleosts (Figure 7), the suprapharyngobranchial

eb
(a) (b) eb (c)
eb

iac

pb pb pb pbtp
pb pb

Figure 7 Dorsal portion of the branchial arches of Amia (a), Hiodon (b), and Trachurus (c) in dorsal view. Cartilage is shown in dark blue,
chondral bone in purple, and dermal bone in light green. Abbreviations: eb, epibranchial; iac, interarcual cartilage; pb, pharyngobranchial;
pbtp, pharyngobranchial toothplate. Modified from Grande L and Bemis WE (1998) A comprehensive phylogenetic study of amiid fishes
(Amiidae) based on comparative skeletal anatomy. An empirical search for interconnected patterns of natural history. Society of Vertebrate
Paleontology, Memoir 4: 1–690; Hilton EJ (2002) Osteology of the extant North American fishes of the genus Hiodon Lesueur, 1818
(Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142; and and Suda Y (1996) Osteology and
muscular attachments of the Japanese jack mackerel, Trachurus japonicus. Bulletin of Marine Science 58: 438–493.
442 The Skeleton | Bony Fish Skeleton

series is absent, and the infrapharyngobranchials (often (basiventrals) found ventrally along the length of the
termed simply ‘pharyngobranchials’) suspend the branchial notochord in early stages of development. The parapo­
basket from the neurocranium. physes are matched by the neural arches, which are
All elements of the branchial arches may support ossifications of the basidorsals (which form on either
toothplates and gill-rakers, which form series on both side of the spinal cord). In basal forms (e.g., sturgeons),
the leading and trailing edges of the arch and function irregularly shaped cartilaginous elements – interdorsals
to keep the gill tissue clear, and they serve to sieve the and interventrals – are intercalated between adjacent
water column to collect food in some filter-feeding taxa basidorsals and basiventrals, respectively. Both neural
(e.g., many Clupeidae). The number and arrangement of arches and parapophyses may fuse to the centra, which
the toothplates, particularly on the dorsal branchial arch typically ossify within the notochordal sheath (although
skeleton, often have significant phylogenetic and func­ there is substantial variation in the mode of ossification
tional significance. The structure of the branchial arches of centra, e.g., some parts of a centrum ossify in conjunc­
generally can inform greatly about the ecology of fishes tion with the basidorsals and basiventrals in addition to
and is rich in phylogenetically informative character data. the notochord). In some basal actinopterygians (e.g.,
sturgeons) and fish-like sarcopterygians (coelacanths
and lungfishes), the centra do not ossify (the so-called
Postcranial Axial Skeleton unconstricted notochord), although the other elements of
the vertebrae are present.
The postcranial axial skeleton of fishes comprises three In actinopterygians, the neural arches extend dorsally,
basic units – the scale jacket, the vertebral skeleton, and forming the neural spine. In derived teleosts, most neural
the median fins and their supporting skeletal elements. and haemal spines are not preformed in cartilage but are
The scales of fishes are widely varied in their structure. membrane bone. The left and right neural spines typically
These range from the large, bony cosmoid scales of most fuse forming a median structure, although the two may
fossil fish-like sarcopterygians, which contain a layer of
remain separate in basal actinopterygians. Intercalated
cosmine (lost in extant sarcopterygians), to the thin, flex­
between the neural spines anterior to the dorsal fin is a
ible cycloid scales found in most teleostean fishes. Some
series of supraneural bones. The centra and neural arches
teleosts have so-called ctenoid scales, which are similar to
of the abdominal region continue into the caudal region
cycloid scales, but which have small spines (cteni) on the
(Figures 8(c) and 8(d)). The parapohyses, on the other
exposed posterior field of the scale. Basal sarcopterygians
hand, end at the abdominal–caudal division and are
and basal actinopterygians (e.g., Polypterus, gars) have
replaced by the haemal arches and spines. Like the para­
thick, interlocking ganoid scales that are coated with
pophyses, these structures develop from the basiventrals,
ganoine (enamel and dentine). Scales (or in some cases
but are similar in form to the neural arches. The hemal
other dermal ossifications) support the lateral sensory
canal (if present) and serve to protect the fish. If the scales arches surround the major blood vessels of the caudal
are reduced in size, there may be other functional region.
protecting layers that are developed (e.g., the greatly Intermuscular bones are associated with the other ele­
thickened dermis of the ocean sunfish, Mola mola). Scales ments of the vertebrae of teleosts, and these bones are
and the dermis itself may also store energy used during widely regarded as a synapomorphy for the Teleostei.
locomotion and respiration. More broadly speaking, the These bones form in the ligaments of the myosepta, and
scale jacket may also include other bony structures, such are thought to help stiffen the body and provide more
as scutes (bony plates serially arranged along the body, as attachment sites for transmission of force. Only relatively
in sturgeons, some catfishes, and carangid fishes) and basal teleosts (e.g., elopomorphs and clupeomorphs) have
fulcra (scales typically associated with the leading edge a complete set of intermuscular bones (i.e., epineurals,
of fins). epicentrals, epipleurals, and myorhabdoi, a naming
The vertebral column can be divided into abdominal based on which fold of the myosepta they are found
and caudal regions, with the caudal region further within), and their presence, absence, and positioning are
divided into the preural and ural regions. The number, phylogenetically informative.
form, and modifications of the vertebrae (e.g., zygapo­ The elements of the vertebral column continue poster­
physes, which may lock together sequential vertebrae in iorly into the ural region (defined as the region posterior
groups such as some billfishes) greatly affect the flex­ to the vertebra that includes the parhypural, which is the
ibility of the body, and therefore have significant last element that has a hemal arch surrounding the blood
functional correlations. The abdominal region is defined vessels; i.e., the first ural vertebrae is associated with
as that region posterior to the skull with vertebrae bear­ hypural 1), which contains most of the supports for the
ing parapophyses (Figures 8(a) and 8(b)). The caudal fin (Figure 9). The caudal fin and its supports are
parapophyses are ossifications of paired cartilages the source of many phylogenetically informative
The Skeleton | Bony Fish Skeleton 443

(a) (b) (c) (d)


ns ns ns
ns

na na na
epn na

c c c
pp c
rb ha
pp
ha
rb

hs
hs

Figure 8 Abdominal (a, b) and caudal (c, d) vertebrae of Amia (a, c) and Hiodon (b, d) in lateral view. Chondral bone is shown in purple,
membrane bone in yellow, and perichordal bone in light blue. Abbreviations: c, centrum; epn, epineural intermuscular bone; ha, hemal
arch; hs, hemal spine; na, neural arch; ns, neural spine; pp, parapophysis; rb, rib. Modified from Grande L and Bemis WE (1998)
A comprehensive phylogenetic study of amiid fishes (Amiidae) based on comparative skeletal anatomy. An empirical search for
interconnected patterns of natural history. Society of Vertebrate Paleontology, Memoir 4: 1–690; and Hilton EJ (2002) Osteology of the
extant North American fishes of the genus Hiodon Lesueur, 1818 (Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology)
new series 100: 1–142.

characters and is perhaps the most-studied region of the Teleostei. In derived teleosts, the uroneurals may fuse
skeleton. to the terminal centrum (which is thought to incorporate
In basal sarcopterygians and nonteleostean actinopter­ preural centrum 1 and all ural centra; Figure 9(c)). There
gyians, the caudal fin is heterocercal, in which the are significant functional consequences to the various
vertebral supports bend slightly dorsally, and the ventral patterns found in the caudal skeletons and fins of fishes,
skeletal supports (the hypurals, the serial homologues of although both heterocercal and homocercal tails appear to
the haemal spines) extend to the very tip of the noto­ function asymmetrically (perhaps through active control
chord. The fin rays are almost entirely hypaxial, and by species with homocercal tails). Therefore, even fishes
produce an internally and externally asymmetrical, with a broad diversity of tail forms all may produce
shark-like caudal fin. Nonteleostean actinopterygians asymmetrical patterns of thrust and lift, which are
(e.g., gars, Amia) have an abbreviated heterocercal caudal matched to ecological and behavioral necessities.
skeleton. This is in contrast to the condition seen in The dorsal and anal fins have a similar overall struc­
teleosts, in which the notochord has a sharp bend at its ture and arrangement (Figure 10). In both fins the
posterior end, causing the hypurals to be directed poster­ skeletal supports for the fin rays, collectively called the
iorly. The fin itself is symmetrical, although the internal pterygiophores, are composed of a series of radial ele­
supporting elements are asymmetrical, forming the so- ments termed the proximal, middle, and distal radials.
called homocercal caudal fin. Most notable in the asym­ The proximal radials are typically the longest of the
metry of the internal skeleton is the presence of series. In most derived teleosts, the proximal and middle
uroneurals, which are the homologs to the neural arches radials are fused, forming the so-called proximal-middle
of ural vertebrae that have become separated from their radial. The distal radials are typically the smallest ele­
vertebrae and modified into stabilizing struts that serve to ments of a pterygiophore, and are embraced by the bases
stiffen the vertebrae of the posterior caudal skeleton. The of the fin rays. In neopterygians (gars, bowfins, and tele­
presence of uroneurals is a key synapomorphy of the osts), the fin rays are about equal in number to the number
444 The Skeleton | Bony Fish Skeleton

(a) ep

napu2
ns
hy5
na

hy1
c

ha

hs phy

ep
ns hy5
(b) un

napu2

na

ha hy1

hs

phy

hy5
ep
(c)
napu1

na

tc
c
hy1+2
ha

hs

hyp
phy
Figure 9 Caudal skeletons of Amia (a), Hiodon (b), and Trachurus (c) in lateral view. Cartilage is shown in dark blue, chondral bone in
purple, perichordal bone in light blue, and dermal bone in green. Abbreviations: ep, epural; ha, hemal arch; hs, hemal spine; hy, hypural;
hyp, hypurapophysis; na, neural arch; napu2, neural arch of preural 2; ns, neural spine; phy, parhypural; tc, terminal centrum; un,
uroneural. Modified from Grande L and Bemis WE (1998) A comprehensive phylogenetic study of amiid fishes (Amiidae) based on
comparative skeletal anatomy. An empirical search for interconnected patterns of natural history. Society of Vertebrate Paleontology,
Memoir 4: 1–690; Hilton EJ (2002) Osteology of the extant North American fishes of the genus Hiodon Lesueur, 1818 (Teleostei:
Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142 and Suda Y (1996) Osteology and muscular
attachments of the Japanese jack mackerel, Trachurus japonicus. Bulletin of Marine Science 58: 438–493.
The Skeleton | Bony Fish Skeleton 445

(a) (b) (d)

(c)
dr
ht(r) ht(l) pmr
sp

mr dr
dr (e)
pr
mr
dr
dr
pr pmr
pmr

(f)
sn

ns

Figure 10 Median fin supports of Amia (a, b) and Trachurus (c–f). (a) Anterior view of a single fin ray and supporting elements.
(b) Lateral view of five pterygiophores; fin rays removed from last three to show distal radials. (c) Oblique view of third dorsal spine.
Elements are separated to show complex structures (dashed lines indicate articulations). (d) Articulated and (e) disarticulated soft fin
rays and their support. (f) Pterygiophores from the spinous and anterior portion of the soft dorsal fin. Cartilage is shown in dark blue,
chondral bone in purple, and dermal bone in green. Abbreviations: dr, distal radial; ht(l), left hemilepidotrichia; ht(r), right
hemilepidotrichia; mr, middle radial; ns, neural spine; pmr, proximal–middle radial; pr, proximal radial; sn, supraneural; sp, spine.
Modified from Grande L and Bemis WE (1998) A comprehensive phylogenetic study of amiid fishes (Amiidae) based on comparative
skeletal anatomy. An empirical search for interconnected patterns of natural history. Society of Vertebrate Paleontology, Memoir
4: 1–690 and Suda Y (1996) Osteology and muscular attachments of the Japanese jack mackerel, Trachurus japonicus. Bulletin of
Marine Science 58: 438–493.

of pterygiophores, which stands in contrast to more the epioccipital) and a ventral limb that contacts or braces
basal actinopterygians (e.g., sturgeons), in which the fin against the skull (often at the intercalar) through ligament
rays greatly outnumber the pterygiophores. The fin rays or direct bony articulation. Ventral to the posttemporal is
(lepidotrichia) of nonacanthomorph teleosts are all soft the supracleithrum, which is attached to the axial skeleton
rays formed by paired hemilepidotrichia, with the excep- (either the basioccipital or an anterior vertebra) through
tion of catfishes and some cyprinids, which have leading the so-called Baudelot’s ligament. The supracleithrum
spine-like fin rays. In Acanthomorpha, a series of true, and the posttemporal carry the trunk lateral sensory
unsegmented median spines precede the soft rays of both canal to the skull. The largest dermal bone of the pectoral
the dorsal and anal fins, and may form a separate spinous girdle is the cleithrum, which is typically a J-shaped bone
dorsal fin. that provides much of the surface area for muscular
attachment in the pectoral girdle. In basal actinoptery­
gians and in sarcopterygians, there is also a clavicle that is
Appendicular Skeleton positioned anteroventral to the cleithrum. The clavicles
are lost in gars and all teleosts. In the bowfin, Amia, there
The pectoral girdle of most actinopterygian fishes are two small plates that are superficial to the sternohyoi­
(Figure 11) is unique among vertebrates in having a deus muscle that have been homologized with the
direct and often strong attachment to the posterior region clavicles of more basal actinopterygians. Also in basal
of the skull. This connection is achieved through the actinopterygians and sarcopterygians, there is a median
posttemporal bone. The posttemporal most often bears dermal interclavicle intercalated between the clavicles;
dorsal limb that contacts the posterodorsal portion of the this bone is lost in Neopterygii. Posteriorly, there is a
skull roof or the braincase (e.g., with the extrascapular or series of postcleithra (often one to three elements, but
446 The Skeleton | Bony Fish Skeleton

(a) (b) (c)


pt pt pt

scl

scl
scl
cl

sc pcl
dr
clv pcl
co
cl pcl
cl
ra

scc
co
mtg

pt
(d) (e)
scl mco
(f)
sc
scl
dr
pcl
pcl co
ra
cl cl mco
sc
ra ra
co
scc
mtg
dr
Figure 11 Pectoral girdle of Amia (a, d), Hiodon (b, e, f), and Trachurus (c) in lateral (a–c) and medial (d–f) views. Cartilage is shown in
dark blue, chondral bone in purple, and dermal bone in green. Abbreviations: cl, cleithrum; clv, clavicle; co, coracoid; dr, distal radial;
mco, mesocoracoid; mtg, metapterygium; pcl, postcleithrum; pt, posttemporal; ra, radial; sc, scapula; scc, scapulocoracoid cartilage;
scl, supracleithrum. Modified from Grande L and Bemis WE (1998) A comprehensive phylogenetic study of amiid fishes (Amiidae) based
on comparative skeletal anatomy. An empirical search for interconnected patterns of natural history. Society of Vertebrate
Paleontology, Memoir 4: 1–690; Hilton EJ (2002) Osteology of the extant North American fishes of the genus Hiodon Lesueur, 1818
(Teleostei: Osteoglossomorpha: Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142; and Suda Y (1996) Osteology and
muscular attachments of the Japanese jack mackerel, Trachurus japonicus. Bulletin of Marine Science 58: 438–493.

sometimes more or less), which extend posteroventrally actinopterygians: the scapula, the coracoid, and the meso­
from the supracleithrum–cleithrum joint. In many teleosts coracoid. The scapula ossifies dorsally, surrounding the
the posteriormost postcleithrum is greatly elongated and scapular foramen, whereas the coracoid ossifies ventrally
rib-like in its form. In sarcopterygians, the shoulder girdle within the scapulocoracoid cartilage. The mesocoracoid
is often separate from the skull (e.g., coelacanths and ossifies in a medial arch of the scapulocoracoid cartilage
lungfishes), but may have additional dermal bones that braces against the medial surface of the cleithrum.
(e.g., the anocleithrum and extracleithrum of coelacanths). The mesocoracoid is lost in advanced teleostean fishes. In
The internal chondral portion of the pectoral girdle of Sarcopterygii, a single element articulates with the
bony fishes is formed by the scapulocoracoid cartilage and scapulocoracoid, forming the so-called monobasal fin.
its ossifications, and the radial elements that directly The proximal radials of the pectoral fin of most actinop­
support the fin rays. In some taxa, there may be additional terygians articulate with the scapula and the coracoid and
neomorphic elements that are synapomorphies for these extend distally to support the fin rays. Typically, there
groups (e.g., the supracleithral cartilage of sturgeons, are four or more proximal radials (=actinosts) that ossify
family Acipenseridae). Three endochondral bones are and a single propterygium that often becomes fused to the
present in nonteleostean and basal teleostean leading fin ray. In nonteleostean actinopterygians, there is
The Skeleton | Bony Fish Skeleton 447

(a) ra teleostean fishes (e.g., Acanthomorpha), the position of


the pelvic fin is shifted forward on the body, and in
many acanthomorphs, the pelvic fins may be anterior to
the pectoral fins. The migration of the pelvic fin has both
mtg phylogenetic (e.g., occurs as a derived state) as well as
plb
functional significance, and seems to be correlated
with increased maneuverability and control during loco­
motion. In Acanthopterygii the soft fin rays of the pelvic
fin are preceded by a stout fin spine.
The largest element of the pelvic girdle of most bony
fishes is the pelvic bone (basipterygium), which forms a
large plate-like element that serves to support the pelvic
fin musculature (Figure 12). In nonteleostean fishes, the
(b) anterior ends of the left and right basipterygia contact or
slightly overlap one another. In sturgeons and paddle­
fishes, the left and right pelvic girdles are widely
pplb
plb separated, each embedded in the hypaxial musculature
of the body wall. In most teleosts, the posterior regions of
the basipterygia are in contact with the midline. Pelvic
ra
radials are lost in the adults of derived acanthomorphs.

Conclusion
(c)
plb
Collectively, the skeletons of fishes display a complex and
diverse array of forms. The details of this variation, which
are merely hinted at in this chapter, are the product of
functional demands generated by the life history, ecology,
Figure 12 Pelvic girdle of Amia (a), Hiodon (b), and Trachurus and behavior of individual species. Coupled to this are the
(c) in dorsal view (inset in (b) shows base of pelvic bone and constraints set in place from the phylogenetic history of
radials with fin rays removed). Cartilage is shown in dark blue, each species. It is this combination – phylogeny and
chondral bone in purple, and dermal bone in green. function – that makes an understanding and appreciation
Abbreviations: mtg, metapterygium; plb, pelvic bone; pplb,
postpelvic bone; ra, radial. Modified from Grande L and Bemis
of the skeleton of fishes central to so many aspects of the
WE (1998) A comprehensive phylogenetic study of amiid fishes evolutionary and ecological biology of fishes.
(Amiidae) based on comparative skeletal anatomy. An empirical
search for interconnected patterns of natural history. Society of See also: The Muscles: Bony Fish Cranial Muscles.
Vertebrate Paleontology, Memoir 4: 1–690; Hilton EJ (2002)
Osteology of the extant North American fishes of the genus
Hiodon Lesueur, 1818 (Teleostei: Osteoglossomorpha:
Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142; and Further Reading
Suda Y (1996) Osteology and muscular attachments of the
Arratia G (2008) Actinopterygian postcranial skeleton with special
Japanese jack mackerel, Trachurus japonicus. Bulletin of Marine
reference to the diversity of fin ray elements and the problem of
Science 58: 438–493. identifying homologies. In: Arratia G, Schultze H-P, and Wilson MVH
(eds.) Mesozoic Fishes 4 – Homology and Phylogeny, pp. 49–101.
München: Verlag Pfeil.
also a posteriorly positioned, elongated metapterygoid Arratia G (2009) Identifying patterns of diversity of the actinopterygian
fulcra. Acta Zoologica (Stockholm) 90(supplement 1): 220–235.
that contributes to the support of the proximal radials. Arratia G and Schultze H-P (1991) Palatoquadrate and its ossifications:
Finally, there is a series of distal radials, which are Development and homology within osteichthyans. Journal of
Morphology 208: 1–81.
entirely surrounded by the bases of the fin rays of the Arratia G and Schultze H-P (1992) Reevaluation of the caudal skeleton
pectoral fin. In sturgeons, there is a pectoral fin spine that of some actinopterygian fishes: III. Salmonidae. Homologization of
is formed by the fusion of adjacent fin rays. In catfishes, caudal skeletal structures. Journal of Morphology 214: 187–249.
Arratia G, Schultze H-P, and Casciotta J (2001) Vertebral column and
the pectoral fin also has a pectoral fin spine formed by the associated elements in dipnoans and comparison with other fishes:
fusion of the segments of the leading fin ray. In these Development and homology. Journal of Morphology 250: 101–172.
spines, the posterior margin may be serrated. Forey PL (1998) History of the Coelacanth Fishes. London: Chapman
and Hall.
The pelvic fin of basal taxa is positioned posteriorly on Grande L and Bemis WE (1998) A comprehensive phylogenetic study of
the ventral portion of the abdominal region. In derived amiid fishes (Amiidae) based on comparative skeletal anatomy. An
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empirical search for interconnected patterns of natural history. Rosen DE, Forey PL, Gardiner BG, and Patterson C (1981) Lungfishes,
Society of Vertebrate Paleontology, Memoir 4: 1–690. tetrapods, paleontology and plesiomorphy. Bulletin of the American
Hilton EJ (2002) Osteology of the extant North American fishes of the Museum of Natural History 167: 159–276.
genus Hiodon Lesueur, 1818 (Teleostei: Osteoglossomorpha: Schultze H-P (1996) The scales of Mesozoic actinopterygians.
Hiodontiformes). Fieldiana (Zoology) new series 100: 1–142. In: Arratia G and Viohl G (eds.) Mesozoic Fishes: Systematics and
Jessen H (1972) Schultergürtel und Pectoralflosse bei Actinopterygiern. Paleoecology, pp. 83–93. München: Verlag Pfeil.
Fossils and Strata 1: 1–101. Schultze H-P (2008) Nomenclature and homologization of cranial bones
Nelson GJ (1969) Gill arches and the phylogeny of fishes, with notes on in actinopterygians. In: Arratia G, Schultze H-P, and Wilson MVH
the classification of vertebrates. Bulletin of the American Museum of (eds.) Mesozoic Fishes 4 – Homology and Phylogeny, pp. 23–48
Natural History 141: 475–552. München: Verlag Pfeil.
Patterson C (1975) The braincase of pholidophorid and leptolepid Schultze H-P and Arratia G (1986) Reevaluation of the caudal skeleton
fishes, with a review of the actinopterygian braincase. Philosophical of some actinopterygian fishes: I. Lepisosteus and Amia. Journal of
Transactions of the Royal Society 269: 275–579. Morphology 190: 215–241.
Patterson C (1977) Cartilage bones, dermal bones, and membrane Schultze H-P and Arratia G (1988) Reevaluation of the caudal skeleton
bones, or the exoskeleton versus the endoskeleton. In: Andrews SM, of some actinopterygian fishes: II. Hiodon, Elops, and Albula. Journal
Miles RS, and Walker AD (eds.) Problems in Vertebrate Evolution. of Morphology 195: 257–303.
pp. 77–121. London: Academic Press. Shadwick RE and Lauder GV (eds.) (2006) Fish Biomechanics. San
Patterson C and Johnson GD (1995) The intermuscular bones and Diego: Elsevier Academic Press.
ligaments of teleostean fishes. Smithsonian Contributions to Zoology Suda Y (1996) Osteology and muscular attachments of the Japanese jack
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