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Helicoprion

Helicoprion is a genus of extinct, shark-


like[2] eugeneodontid holocephalid fish.
Almost all fossil specimens are of
spirally arranged clusters of the
individuals' teeth, called "tooth whorls"—
the cartilaginous skull, spine, and other
structural elements have not been
preserved in the fossil record, leaving
scientists to make educated guesses as
to its anatomy and behavior. Helicoprion
lived in the oceans of the early Permian[3]
290 million years ago, with species
known from North America, Eastern
Europe, Asia, and Australia.[4] The closest
living relatives of Helicoprion (and other
eugeneodontids) are the chimaeras.[5]
Helicoprion
Temporal range: Cisuralian-Guadalupian
PreЄ Є O S D C P T J K PN
g

Tooth-whorl, Utah Field House of Natural


History

Scientific classification

Kingdom: Animalia

Phylum: Chordata

Class: Chondrichthyes

Order: †Eugeneodontida

Family: †Helicoprionidae

Genus: †Helicoprion
Karpinsky, 1899[1]

Type species

Helicoprion bessonowi
Karpinsky, 1899

Species

H. bessonowi Karpinsky, 1899


H. davisii Woodward, 1886
H. ergasaminon Bendix-Almgreen,
1966
?H. karpinskii Obruchev, 1953
?H. mexicanus Mullerried, 1945
?H. svalis Siedlecki, 1970

Description
Illustration of H. bessonowi

In 2011, a tooth whorl from a Helicoprion


was discovered in the Phosphoria site in
Idaho. The tooth whorl measured 45 cm
(18 in) in length. Comparisons with other
Helicoprion specimens show that the
animal that sported this whorl would
have been 10 m (33 ft) in length, and
another, even bigger tooth whorl that was
discovered in 1980s (but was not
published until 2013) which the
discoverers dubbed IMNH 49382 or
"Boise" was discovered at the same site.
The whorl is incomplete, but in life it
would have been 60 cm (24 in) long and
would have belonged to an animal that
possibly exceeded 12 m (39 ft) in length,
making Helicoprion the largest known
eugeneodont.[6]

Tooth-whorl

Alexander Karpinsky's 1899 hypothesis of the


placement of the tooth whorl on H. bessonowi.
Outdated illustration of H. bessonowi, showing
tooth-whorl at the front of the jaw

Until 2013, the only known fossils of this


genus on record were their teeth, which
were arranged in a "tooth-whorl" strongly
reminiscent of a circular saw. As the
skeletons of chondrichthyid fish are
made of cartilage, including those of
Helicoprion and other eugeneodonts, the
entire body disintegrates once it begins
to decay, unless exceptional
circumstances preserve it. The tooth-
whorl was not realized to be in the lower
jaw until the discovery of the skull of a
related genus of eugeneodont,
Ornithoprion. The tooth-whorl
represented all the teeth produced by
that individual in the lower jaw; as the
individual grew, the older, smaller teeth
were moved into the center of the whorl
by larger, newer teeth appearing. Models
of the Helicoprion tooth-whorl have been
made. In the 1994 book Planet Ocean: A
Story of Life, the Sea, and Dancing to the
Fossil Record, author Brad Matsen and
artist Ray Troll describe and depict an
example of such a model. They proposed
that no teeth were present in the animal's
top jaw besides the crushing teeth for the
whorl to cut against. The two envision
the living animal to have a long and very
narrow skull, creating a long nose akin to
the modern-day goblin shark. According
to their studies, the fossils that have
been found are essentially a growth ring,
as each set of new teeth pushes the
previous set into the whorl.[7]

For over a century, whether the tooth-


whorl was situated in the lower jaw
wasn't certain. Older reconstructions
placed the whorl in the front of the lower
jaw. A 2008 reconstruction, created by
Mary Parrish under the direction of
Robert Purdy, Victor Springer, and Matt
Carrano for the Smithsonian, places the
whorl deeper into the throat,[3] although
other studies did not accept this
conclusion.[8][9] A 2013 study based on
new data places the tooth-whorl at the
back of the jaw, where the tooth-whorl
occupied the entire mandibular arch.[5]

Cast of tooth-whorl

In his 1939 article, author Harry E.


Wheeler describes another Helicoprion
fossil, based on the species H. sierrensis,
collected by J. H. Menke, that resides at
the University of Nevada, W. M. Keck
Earth Science and Mineral Engineering
Museum. This fossil is number 1002 and
is currently on display in case 62. The
mouth consists of a whorl separated into
three and a quarter volutions. The
biggest diameter is about 170 mm
(6.7 in). The whorls have a separation of
about 1 mm (0.039 in) in the first
volution, and it goes to about 8 mm
(0.31 in) at the largest whorl displayed.
The specimen has a total of about 32
teeth in the first volution, 36 in the
second, and 41 in the last. The teeth at
the end of the first volution are about
7 mm (0.28 in) long and are about 2.4 in
(61 mm) width reaching about 40 mm
(1.6 in) long and 9.5 wide at the end of
the third. The teeth are symmetrically
opposed to one another.[10]

Additionally, other extinct fish, such as


Onychodontiformes, have analogous
tooth-whorls at the front of the jaw,
suggesting that such whorls are not as
big of an impediment to swimming as
suggested in Purdy's hypothesis. While
no complete skulls of Helicoprion have
been officially described, the fact that
related species of chondrichthyids had
long, pointed snouts suggests that
Helicoprion did, as well.

Distribution
Helicoprion species proliferated greatly
during the early Permian. Fossils have
been found in the Ural Mountains,
Western Australia, China[11] (together
with the related genera Sinohelicoprion
and Hunanohelicoprion), and Western
North America, including the Canadian
Arctic, Mexico, Idaho, Nevada, Wyoming,
Texas, Utah, and California. More than
50% of Helicoprion specimens are known
from Idaho, with an additional 25% being
found in the Ural Mountains.[4] Due to the
fossils' locations, the various species of
Helicoprion may have lived off the
southwestern coast of Gondwana, and
later, Pangaea.
Species
H. bessonowi

Restoration of H. bessonowi

Helicoprion was first described by


Alexander Karpinsky in 1899 from a
fossil found in Artinskian age limestones
of the Ural Mountains.[12] Karpinsky
named the type species Helicoprion
bessonowi; Oliver Perry Hay originally
described the species. This species can
be differentiated from others by a short
and narrowly spaced tooth whorl,
backward-directed tooth tips, obtusely-
angled tooth bases, and a consistently
narrow whorl shaft.[4]

One of two Helicoprion species described


by Wheeler in 1939, H. nevadensis, is
based on a single partial fossil found in
1929 by Elbert A Stuart.[10] It was
reported as having originated from the
Rochester Trachyte deposits, which
Wheeler considered to be of Artinskian
age. However, the Rochester Trachyte is
in fact Triassic, and H. nevadensis likely
did not originate in the Rochester
Trachyte, thus rendering its true age
unknown. Wheeler differentiated H.
nevadensis from H. bessonowi by its
pattern of whorl expansion and tooth
height, but Leif Tapanila and Jesse Pruitt
showed in 2013 that these were
consistent with H. bessonowi at the
developmental stage that the specimen
represents.[4]

Based on isolated teeth and partial


whorls found on the island of
Spitsbergen, Norway, H. svalis was
described by Stanisław Siedlecki in 1970.
The type specimen, a very large whorl,
was noted for its narrow teeth that
apparently are not in contact with each
other. However, this seems to be a
consequence of only the central part of
the teeth being preserved, according to
Tapanila and Pruitt. Since the whorl shaft
is partially obscured, H. svalis cannot be
definitely assigned to H. bessonowi, but it
closely approaches the latter species in
many aspects of its proportions. With a
maximum volution height of 72
millimetres (2.8 in), H. svalis is similar in
size to the largest H. bessonowi, which
has a maximum volution height of 76
millimetres (3.0 in).[4]

H. davisii

H. davisii was described initially from a


series of 15 teeth found in Western
Australia. They were described by H.
Woodward in 1886 as a species of
Edestus, E. davisii. Upon naming H.
bessonowi, Karpinsky also reassigned
this species to Helicoprion, an
identification subsequently supported by
the discovery of two additional and more
complete tooth whorls in Western
Australia. The species is characterized by
a tall and widely spaced tooth whorl, with
these becoming more pronounced with
age. The teeth also noticeably curve
forwards. During the Kungurian and
Roadian, this species was very common
worldwide.[4]
H. davisii (=H. ferrieri) tooth-whorl, Glass Mountains,
Texas

H. ferrieri was originally described as a


species of the genus Lissoprion in 1907,
from fossils found in the Phosphoria
Formation of Idaho. An additional
specimen, tentatively referred to H.
ferrieri, was described in 1955. That
specimen was found in Wolfcampian-age
quartzites exposed on China Mountain,
six miles southeast of Contact, Nevada.
The 100-mm-wide fossil consists of one
and three-quarters whorls and about 61
preserved teeth. Due to weathering, the
rest of the fossil was lost and the
preserved section is distorted from
slippage of the host rock.[12] While
initially differentiated using the metrics
of tooth angle and height, Tapanila and
Pruitt considered these characteristics to
be intraspecifically variable, reassigning
H. ferrieri to H. davisii.[4]

H. jingmenense was described in 2007


from a nearly complete tooth whorl with
four and a third volutions (part and
counterpart) found in the Lower Permian
Qixia Formation of Hubei Province,
China. It was discovered during road
construction. The specimen is very
similar to H. ferrieri and H. bessonowi,
though it differs from the former by
having teeth with a wider cutting blade,
and a shorter compound root, and differs
from the latter by having fewer than 39
teeth per volution.[11] Tapanila and Pruitt
argued that the specimen was partially
obscured by the surrounding matrix,
resulting in an underestimation of tooth
height. Taking into account intraspecific
variation, they synonymized it with H.
davisii.[4]

H. ergassaminon

H. ergassaminon, the rarer species from


the Phosphoria Formation, was
described in detail within a 1966
monograph by Svend Erik Bendix-
Almgreen. The holotype specimen
("Idaho 5"), now lost, bore breakage and
wear marks indicative of its usage in
feeding. Several referred specimens
exist, none of which show wear marks.
This species is roughly intermediate
between the two contrasting forms
represented by H. bessonowi and H.
davisii, having tall but narrowly-spaced
teeth. Its teeth are also gently curved,
with obtusely-angled tooth bases.[4]

Other material
Several large whorls are difficult to
assign to any particular species group, H.
svalis among them. IMNH 14095, a
specimen from Idaho, appears to be
similar to H. bessonowi, but it has unique
flange-like edges on the apices of its
teeth. IMNH 49382, also from Idaho, has
the largest known whorl diameter at 56
millimetres (2.2 in) for the outermost
volution (the only one preserved), but it is
incompletely preserved and still partially
buried.[4]

H. mexicanus, named by F.K.G. Müllerreid


in 1945 and supposedly distinguished by
its tooth ornamentation, has a holotype
that is currently missing, but its
morphology was similar to IMNH 49382.
In the absence of other material, it is
currently a nomen dubium. Vladimir
Obruchev described H. karpinskii from
two teeth in 1953. He provided no
distinguishing traits for this species, and
thus it must be regarded as a nomen
nudum.[4]

References
1. Карпинскій, А. (1899). Объ
остаткахъ eдестидъ и о новомъ
ихъ родѣ Helicoprion [On the
edestid remains and the new genus
Helicoprion]. Записки
Императорской Академіи Наукъ
(Notes of the Imperial Academy of
Sciences). По Физико-
математическому отдѣленіи
(Physics and Mathematics section)
(in Russian). 8 (7): 1–67; Pl. I–IV.
Also printed as "Ueber die Reste von
Edestiden und die neue Gattung
Helicoprion" . Записки
Императорскаго С.-
Петербургскаго
Минералогическаго
Обществ(Notes of the Imperial St.
Petersburg Mineralogical Society). 2
(in German). 36: 361–476. 1899; Pl.
I–IV. [Reprinted (1899). St.
Petersburg: C. Birkenfield. pp. 1–
111. urn:nbn:de:bvb:12-
bsb00073910-2 .]
2. Viegas, Jennifer (February 27, 2013).
"Ancient shark relative had buzzsaw
mouth" . science.nbcnews.com.
3. Purdy, Robert (February 29, 2008).
"The Orthodonty of Helicoprion" .
paleobiology.si.edu. Smithsonian.
Archived from the original on June
18, 2018.
4. Tapanila, L.; Pruitt, J. (2013).
"Unravelling species concepts for the
Helicoprion tooth whorl" (PDF).
Journal of Paleontology. 87 (6):
965–983. doi:10.1666/12-156 .
5. Tapanila, L.; Pruitt, J.; Pradel, A.;
Wilga, C.D.; Ramsay, J.B.; Schlader,
R.; Didier, D.A. (2013). "Jaws for a
spiral-tooth whorl: CT images reveal
novel adaptation and phylogeny in
fossil Helicoprion" . Biology Letters.
9 (2): 20130057.
doi:10.1098/rsbl.2013.0057 .
PMC 3639784 . PMID 23445952 .
6. Udurawane, Vasika. "Buzzsaw-
toothed leviathans cruised the
ancient seas" . eartharchives.org.
7. Matsen, Brad; Troll, Ray (1994).
Planet Ocean: A Story of Life, the
Sea, and Dancing to the Fossil
Record . Berkeley, CA: Ten Speed
Press. ISBN 9780898157789.
8. Lebedev, O.A. (2009). "A new
specimen of Helicoprion Karpinsky,
1899 from Kazakhstanian Cisurals
and a new reconstruction of its tooth
whorl position and function". Acta
Zoologica. 90: 171–182.
doi:10.1111/j.1463-
6395.2008.00353.x . ISSN 0001-
7272 .
9. Brian Switek (February 29, 2008).
"Unraveling the Nature of the Whorl-
Toothed Shark" . Laelaps. Wired.
p. 3. Retrieved September 23, 2012.
10. Wheeler, H. E. (1939). "Helicoprion in
the Anthracolithic (Late Paleozoic)
of Nevada and California, and its
Stratigraphic Significance". Journal
of Paleontology. 13 (1): 103–114.
JSTOR 1298628 .
11. Chen, Xiao Hong; Long, Cheng; Yin,
Kai Guo (August 2007). "The first
record of Helicoprion Karpinsky
(Helicoprionidae) from China".
Chinese Science Bulletin. 52 (16):
2246–2251. doi:10.1007/s11434-
007-0321-y .
12. Larson, E. R.; Scott, J. B. (1955).
"Helicoprion from Elko County,
Nevada". Journal of Paleontology. 29
(5): 918–919. JSTOR 1300414 .

Further reading
Hay, Oliver Perry (1909). "On the nature of
Edestus and related genera, with
descriptions of one new genus and three
new species" . Proceedings of the United
States National Museum. 37 (1699): 43–61.
doi:10.5479/si.00963801.37-1699.43 ; Pl.
12–15.
Карпинскій, А. (1911). "Замѣчанія о
Helicoprion и о другихъ едестидахъ" .
Извѣстія Императорской Академіи Наук =
Bulletin de l'Académie Impériale des
Sciences de St.-Pétersbourg. VI Серия = VI
Série. 5 (16): 1105–1122.
Teichert, Curt (1940). "Helicoprion in the
Permian of Western Australia". Journal of
Paleontology. 14 (2): 140–149.
JSTOR 1298567 .
Woodward, Henry (1886). "On a Remarkable
Ichthyodorulite from the Carboniferous
Series, Gascoyne, Western Australia" .
Geological Magazine. New Series. Decade
III. 3 (1): 1–7.
doi:10.1017/S0016756800144450 .
Yabe, H. (1903). "On a Fusulina-Limeston
with Helicoprion in Japan". The Journal of
the Geological Society of Japan. 10 (113):
1–13. doi:10.5575/geosoc.10.113_1 .

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Last edited on 9 September 2019, at 00:22

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