Chondrichthyes: Skeleton
Chondrichthyes: Skeleton
Chondrichthyes
Chondrichthyes (/kɒnˈdrɪkθiiːz/; from Greek χονδρ- chondr- 'cartilage', ἰχθύς ichthys 'fish') is
a class that contains the cartilaginous fishes: they are jawed vertebrates with paired fins, Cartilaginous fishes
paired nares, scales, a heart with its chambers in series, and skeletons made of cartilage rather Temporal range: 430–0 Ma[1][2]
than bone. The class is divided into two subclasses: Elasmobranchii (sharks, rays, skates, and
PreꞒ Ꞓ O S D C P T J K PgN
sawfish) and Holocephali (chimaeras, sometimes called ghost sharks, which are sometimes
Late Silurian to Present
separated into their own class).
Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed
vertebrates.
Contents
Anatomy
Skeleton Example of cartilaginous fishes : at
Appendages the top of the image, Elasmobranchii
Body covering and at the bottom of the image,
Respiratory system
Holocephali.
Nervous system
Immune system Scientific classification
Subclass Elasmobranchii
Skeleton
Superorder
The skeleton is cartilaginous. The notochord is gradually replaced by a vertebral column during
Selachimorpha
development, except in Holocephali, where the notochord stays intact. In some deepwater
sharks, the column is reduced.[3] Order †Mongolepidida
As they do not have bone marrow, red blood cells are produced in the spleen and the epigonal Order
organ (special tissue around the gonads, which is also thought to play a role in the immune Carcharhiniformes
system). They are also produced in the Leydig's organ, which is only found in certain
cartilaginous fishes. The subclass Holocephali, which is a very specialized group, lacks both the Order Lamniformes
Leydig's and epigonal organs. Order
Orectolobiformes
Appendages Order
Heterodontiformes
Apart from electric rays, which have a thick and flabby body, with soft, loose skin,
chondrichthyans have tough skin covered with dermal teeth (again, Holocephali is an exception, Order Squaliformes
as the teeth are lost in adults, only kept on the clasping organ seen on the caudal ventral surface Order Squatiniformes
of the male), also called placoid scales (or dermal denticles), making it feel like sandpaper. In
most species, all dermal denticles are oriented in one direction, making the skin feel very smooth Order
if rubbed in one direction and very rough if rubbed in the other. Pristiophoriformes
Originally, the pectoral and pelvic girdles, which do not contain any dermal elements, did not Order Hexanchiformes
connect. In later forms, each pair of fins became ventrally connected in the middle when Superorder Batoidea
scapulocoracoid and puboischiadic bars evolved. In rays, the pectoral fins are connected to the
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head and are very flexible. Order Myliobatiformes
One of the primary characteristics present in most sharks is the heterocercal tail, which aids in Order Rajiformes
locomotion.[4] Order
Rhinopristiformes
Body covering Order Torpediniformes
Chondrichthyans have toothlike scales called dermal denticles or placoid scales. Denticles usually Subclass Holocephali
provide protection, and in most cases, streamlining. Mucous glands exist in some species, as well.
Superorder
It is assumed that their oral teeth evolved from dermal denticles that migrated into the mouth, Holocephalimorpha
but it could be the other way around, as the teleost bony fish Denticeps clupeoides has most of its
head covered by dermal teeth (as does, probably, Atherion elymus, another bony fish). This is Order Chimaeriformes
most likely a secondary evolved characteristic, which means there is not necessarily a connection
between the teeth and the original dermal scales.
The old placoderms did not have teeth at all, but had sharp bony plates in their mouth. Thus, it is unknown whether the dermal or oral
teeth evolved first. It has even been suggested that the original bony plates of all vertebrates are now gone and that the present scales
are just modified teeth, even if both the teeth and body armor had a common origin a long time ago. However, there is currently no
evidence of this.
Respiratory system
All chondrichthyans breathe through five to seven pairs of gills, depending on the species. In general, pelagic species must keep
swimming to keep oxygenated water moving through their gills, whilst demersal species can actively pump water in through their
spiracles and out through their gills. However, this is only a general rule and many species differ.
A spiracle is a small hole found behind each eye. These can be tiny and circular, such as found on the nurse shark (Ginglymostoma
cirratum), to extended and slit-like, such as found on the wobbegongs (Orectolobidae). Many larger, pelagic species, such as the
mackerel sharks (Lamnidae) and the thresher sharks (Alopiidae), no longer possess them.
Nervous system
In chondrichthyans, the nervous system is composed of a small brain, 8-10 pairs of cranial nerves,
and a spinal chord with spinal nerves.[5] They have several sensory organs which provide
information to be processed. Ampullae of Lorenzini are a network of small jelly filled pores called
electroreceptors which help the fish sense electric fields in water. This aids in finding prey,
navigation, and sensing temperature. The Lateral line system has modified epithelial cells located
externally which sense motion, vibration, and pressure in the water around them. Most species
have large well-developed eyes. Also, they have very powerful nostrils and olfactory organs. Their
inner ears consist of 3 large semicircular canals which aid in balance and orientation. Their sound
detecting apparatus has limited range and is typically more powerful at lower frequencies. Some
species have electric organs which can be used for defense and predation. They have relatively Regions of a Chondrichthyes brain
simple brains with the forebrain not greatly enlarged. The structure and formation of myelin in colored and labeled on dissected
their nervous systems are nearly identical to that of tetrapods, which has led evolutionary skate. The rostral end of the skate is
biologists to believe that Chondrichthyes were a cornerstone group in the evolutionary timeline of to the right.
myelin development.[6]
Immune system
Like all other jawed vertebrates, members of Chondrichthyes have an adaptive immune system.[7]
Reproduction
Fertilization is internal. Development is usually live birth (ovoviviparous species) but can be through eggs (oviparous). Some rare
species are viviparous. There is no parental care after birth; however, some chondrichthyans do guard their eggs.
Capture-induced premature birth and abortion (collectively called capture-induced parturition) occurs frequently in sharks/rays when
fished.[8] Capture-induced parturition is often mistaken for natural birth by recreational fishers and is rarely considered in commercial
fisheries management despite being shown to occur in at least 12% of live bearing sharks and rays (88 species to date).[8]
Classification
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The class Chondrichthyes has two subclasses: the subclass Elasmobranchii (sharks, rays, skates, and sawfish) and the subclass
Holocephali (chimaeras). To see the full list of the species, click here.
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ground Compagno,
Carcharhiniformes 8 51 >270 7 10 21
sharks 1977
bullhead L. S. Berg,
Heterodontiformes 1 1 9
sharks 1940
Galean
sharks
mackerel L. S. Berg, 7
Lamniformes 10 16 10
sharks 1958 +2 extinct
carpet Applegate,
Orectolobiformes 7 13 43 7
sharks 1972
frilled
de Buen, 2 4 7
Hexanchiformes and
1926 +3 extinct +11 extinct +33 extinct
cow sharks
L. S. Berg,
Pristiophoriformes sawsharks 1 2 6
1958
Squalomorph
sharks
dogfish Goodrich,
Squaliformes 7 23 126 1 6
sharks 1909
angel
Squatiniformes Buen, 1926 1 1 24 3 4 5
sharks
stingrays
Compagno,
Myliobatiformes and 10 29 223 1 16 33
1973
relatives
Rays
skates
L. S. Berg,
Rajiformes and 5 36 >270 4 12 26
1940
guitarfishes
electric de Buen,
Torpediniformes 2 12 69 2 9
rays 1926
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Holocephali Chimaeriformes chimaera Obruchev, 3 6 39
1953 +2 extinct +3 extinct +17 extinct
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†Superorder Paraselachimorpha
†Plesioselachus
†Order Antarctilamniformes Ginter, Liao & Valenzuela-Rios, 2008
†Order Elegestolepidiformes Andreev et al., 2016
†Order Lugalepidida Karatajute-Talimaa, 1997
†Order Squatinactiformes Zangerl, 1981
†Order Protacrodontiformes Zangerl, 1981
†Infraclass Cladoselachimorpha
Order Chlamydoselachiformes
Order Hexanchiformes (frilled and cow sharks)
Order Squaliformes (dogfish sharks)
†Order Protospinaciformes
†Order Synechodontiformes
Order Squatiniformes (angel sharks)
Order Pristiophoriformes (sawsharks)
Subdivision Batoidea
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Order Torpediniformes (electric rays)
Order Pristiformes (sawfishes)
Order Rajiformes (skates and guitarfishes)
Order Myliobatiformes (stingrays and relatives)
* position uncertain
Evolution
Cartilaginous fish are considered to have evolved from acanthodians.
Originally assumed to be closely related to bony fish or a polyphyletic
assemblage leading to both groups, the discovery of Entelognathus and
several examinations of acanthodian characteristics indicate that bony
fish evolved directly from placoderm like ancestors, while acanthodians
represent a paraphyletic assemblage leading to Chondrichthyes. Some
characteristics previously thought to be exclusive to acanthodians are also
present in basal cartilaginous fish.[13] In particular, new phylogenetic
studies find cartilaginous fish to be well nested among acanthodians, with
Doliodus and Tamiobatis being the closest relatives to
Chondrichthyes.[14] Recent studies vindicate this, as Doliodus had a
mosaic of chondrichthyian and acanthodiian traits.[15]
Dating back to the Middle and Late Ordovician Period, many isolated
scales, made of dentine and bone, have a structure and growth form that
is chondrichthyan-like. They may be the remains of stem-
chondrichthyans, but their classification remains uncertain.[16][17][18]
By the start of the Early Devonian, 419 million years ago, jawed fishes had divided into three distinct groups: the now extinct
placoderms (a paraphyletic assemblage of ancient armoured fishes), the bony fishes, and the clade that includes spiny sharks and early
cartilaginous fish. The modern bony fishes, class Osteichthyes, appeared in the late Silurian or early Devonian, about 416 million years
ago. The first abundant genus of shark, Cladoselache, appeared in the oceans during the Devonian Period. The first Cartilaginous fishes
evolved from Doliodus-like spiny shark ancestors.
A Bayesian analysis of molecular data suggests that the Holocephali and Elasmoblanchii diverged in the Silurian
(421 million years ago) and that the sharks and rays/skates split in the Carboniferous (306 million years ago).
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Cladoselache was the first abundant genus of primitive shark, appearing about 370 Ma.[20] It
grew to 6 feet (1.8 m) long, with anatomical features similar to modern mackerel sharks. It had
a streamlined body almost entirely devoid of scales, with five to seven gill slits and a short,
rounded snout that had a terminal mouth opening at the front of the skull.[20] It had a very weak
Devonian jaw joint compared with modern-day sharks, but it compensated for that with very strong jaw-
Cladoselache closing muscles. Its teeth were multi-cusped and smooth-edged, making them suitable for
grasping, but not tearing or chewing. Cladoselache therefore probably seized prey by the tail
and swallowed it whole.[20] It had powerful keels that extended onto the side of the tail stalk and
a semi-lunate tail fin, with the superior lobe about the same size as the inferior. This
combination helped with its speed and agility which was useful when trying to outswim its
probable predator, the heavily armoured 10 metres (33 ft) long placoderm fish Dunkleosteus.[20]
Carboniferous (359–299 Ma): Sharks underwent a major evolutionary radiation during the Carboniferous.[21] It is believed that this evolutionary
radiation occurred because the decline of the placoderms at the end of the Devonian period caused many environmental niches to become
unoccupied and allowed new organisms to evolve and fill these niches.[21]
The first 15 million years of the Carboniferous has very few terrestrial fossils. This gap in the
fossil record, is called Romer's gap after the American palaentologist Alfred Romer. While it has
long been debated whether the gap is a result of fossilisation or relates to an actual event,
recent work indicates that the gap period saw a drop in atmospheric oxygen levels, indicating
some sort of ecological collapse.[22] The gap saw the demise of the Devonian fish-like
ichthyostegalian labyrinthodonts, and the rise of the more advanced temnospondyl and
reptiliomorphan amphibians that so typify the Carboniferous terrestrial vertebrate fauna.
Most species of Carboniferous marine fish have been described largely from
Carbon- teeth, fin spines and dermal ossicles, with smaller freshwater fish preserved
iferous whole. Freshwater fish were abundant, and include the genera Ctenodus,
Uronemus, Acanthodes, Cheirodus, and Gyracanthus.
Falcatus is a genus of small cladodont-toothed sharks which lived 335–318 Ma. They were
Falcatus about 25–30 cm (9.8–11.8 in) long.[23] They are characterised by the prominent fin spines that
curved anteriorly over their heads.
Orodus is another shark of the Carboniferous, a genus from the family Orodontidae that lived
Orodus
into the early Permian from 303 to 295 Ma. It grew to 2 m (6.6 ft) in length.
Permian (298–252 Ma): The Permian ended with the most extensive extinction event recorded in paleontology: the Permian-Triassic extinction
Permian event. 90% to 95% of marine species became extinct, as well as 70% of all land organisms. Recovery from the Permian-Triassic extinction
event was protracted; land ecosystems took 30M years to recover,[24] and marine ecosystems took even longer.[25]
Triassic (252–201 Ma): The fish fauna of the Triassic was remarkably uniform, reflecting the fact that very few families survived the Permian
Triassic extinction. In turn, the Triassic ended with the Triassic–Jurassic extinction event. About 23% of all families, 48% of all genera (20% of marine
families and 55% of marine genera) and 70% to 75% of all species became extinct.[26]
Jurassic Jurassic (201–145 Ma):
Cretaceous Cretaceous (145–66 Ma): The end of the Cretaceous was marked by the Cretaceous–Paleogene extinction event (K-Pg extinction). There are
substantial fossil records of jawed fishes across the K–T boundary, which provides good evidence of extinction patterns of these classes of
marine vertebrates. Within cartilaginous fish, approximately 80% of the sharks, rays, and skates families survived the extinction event,[27] and
more than 90% of teleost fish (bony fish) families survived.[28]
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Squalicorax Squalicorax falcatus is a lamnoid shark from the Cretaceous
falcatus
Ptychodus is a genus of extinct hybodontiform shark which lived from the late Cretaceous to the
Ptychodus Paleogene.[29][30] Ptychodus mortoni (pictured) was about 32 feet (9.8 meters) long and was
unearthed in Kansas, United States.[31]
Cenozoic Era (65 Ma to present): The current era has seen great diversification of bony fishes.
External video
Megalodon is an extinct species of shark Megalodon battle (https://www.youtube.
that lived about 28 to 1.5 Ma. It looked com/watch?v=_9LAWve5Xq4) History
much like a stocky version of the great Channel
Cenozoic white shark, but was much larger with The Nightmarish Megalodon (https://ww
Era Megalodon fossil lengths reaching 20.3 metres w.youtube.com/watch?v=Spo8vkrJFRo)
(67 ft).[32] Found in all oceans[33] it was Discovery
one of the largest and most powerful
predators in vertebrate history,[32] and
probably had a profound impact on marine life.[34]
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†Petalodontiformes
†Helodontiformes
†Iniopterygiformes
†Debeeriiformes
[35]
Holocephali †Symmoriida
[36]
†Eugeneodonti
formes
†Squatinactiformes
†Protacrodonti-
formes
†Cladoselachi-
formes
†Xenacanthiformes
†Ctenacanthi-
formes
†Hybodontiformes
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Taxonomy
Subphylum Vertebrata
└─Infraphylum Gnathostomata
├─Placodermi — extinct (armored gnathostomes)
└Eugnathostomata (true jawed vertebrates)
├─Acanthodii (stem cartilaginous fish)
└─Chondrichthyes (true cartilaginous fish)
├─Holocephali (chimaeras + several extinct clades)
└Elasmobranchii (shark and rays)
├─Selachii (true sharks)
└─Batoidea (rays and relatives)
See also
List of cartilaginous fish
Cartilaginous versus bony fishes
Largest cartilaginous fishes
Threatened rays
Threatened sharks
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Further reading
Taxonomy of Chondrichthyes (https://web.archive.org/web/20040817003309/http://www.fmnh.helsinki.fi/users/haaramo/Metazoa/D
euterostoma/Chordata/Chondrichthyes/Chondrichthyes.htm#Elasmobranchii)
Images of many sharks, skates and rays on Morphbank (http://www.morphbank.net/Browse/ByImage/?tsn=159785)
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