Ischyrodon
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''Ischyrodon'' (meaning 'strong tooth') is a dubious
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of large
pliosaurid Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
plesiosaur The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
from the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
of Switzerland. The only known
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
is ''I. meriani'', originally mentioned in 1838 by
Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born in Frankfurt am ...
from a large tooth discovered in Wölflinswil, before being first described in detail by the same author in 1856. The
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
remained valid until 1889, when
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...
synonymized it with a large species of ''
Pliosaurus ''Pliosaurus'' (meaning 'more lizard') is an extinct genus of thalassophonean pliosaurid known from the Late Jurassic (Kimmeridgian and Tithonian stages) of Europe and South America. This genus has contained many species in the past but recent ...
''. However, in 1960, Lambert Beverly Tarlo noted that the tooth shares more similarities with those of ''
Liopleurodon ''Liopleurodon'' (; meaning 'smooth-sided teeth') is an extinct genus of carnivorous pliosaurid pliosaurs that lived from the Callovian stage of the Middle Jurassic to the Kimmeridgian stage of the Late Jurassic period (c. 166 to 155 mya). T ...
'', instead considering it as a
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of this latter, a view that was later largely followed in
scientific literature Scientific literature encompasses a vast body of academic papers that spans various disciplines within the natural and social sciences. It primarily consists of academic papers that present original empirical research and theoretical ...
. However, a 2022 revision found few differences that would allow it to be distinguished from ''Liopleurodon'' and other
thalassophonea Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
n pliosaurids, then considering it as a ''nomen dubium''.


Research history

The only known specimen of ''Ischyrodon'' is a large tooth that was discovered in Wölflinswil, in the
Canton of Aargau Aargau ( ; ), more formally the Canton of Aargau (; ; ; ), is one of the 26 cantons forming the Swiss Confederation. It is composed of eleven districts and its capital is Aarau. Aargau is one of the most northerly cantons of Switzerland, by th ...
, Switzerland. More precisely, the tooth was probably discovered in an
iron mine Iron is a chemical element; it has symbol Fe () and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, forming much of Earth's ou ...
containing rocks dating from the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
, exploited from the 1200s until 1967. The
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
, consisting precisely of a
tooth crown In dentistry, the crown is the visible part of the tooth above the gingival margin and is an essential component of dental anatomy. Covered by Tooth enamel, enamel, the crown plays a crucial role in cutting, tearing, and grinding food. Its shap ...
measuring almost high, has since been preserved at the
Natural History Museum of Basel Natural History Museum Basel () is a natural history museum in Basel, Switzerland that houses wide-ranging collections focused on the fields of zoology, entomology, mineralogy, anthropology, osteology and paleontology. It has over 11 million obje ...
, from where it is numbered as NMB L.D.37. This tooth is directly recognized by the Swiss naturalist Peter Merian as coming from a new
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
of a large
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. Only about 100 of the 12,000 extant reptile species and subspecies are classed as marine reptiles, including mari ...
, from which he named it ''Ischyrodon'', although he never published a paper officially naming it. The
genus name Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial spec ...
comes from the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
(, "strong") and (', "tooth"), all meaning "strong tooth", because of its large size. In
scientific literature Scientific literature encompasses a vast body of academic papers that spans various disciplines within the natural and social sciences. It primarily consists of academic papers that present original empirical research and theoretical ...
, this name is first mentioned by the German paleontologist
Christian Erich Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born in Frankfurt am ...
in 1838, in a conversation informing the editors of the existence of fossil specimens that had been recently discovered at that time, but without any descriptions or figures. Still in the same conversation, the author named it ''Ischyrodon meriani'', in honor of Merian. Although mentioned in several articles and manuals over the following years, this tooth crown was not first formally described and illustrated until 1856, still by von Meyer, 18 years after the publication of its
scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
. In 1841, von Meyer named the species '' Thaumatosaurus oolithicus'' based on a fragmentary specimen consisting of partial teeth, skull elements, vertebrae, and ribs from deposits in
Württemberg Württemberg ( ; ) is a historical German territory roughly corresponding to the cultural and linguistic region of Swabia. The main town of the region is Stuttgart. Together with Baden and Province of Hohenzollern, Hohenzollern, two other histo ...
, Germany, possibly dating to the Oxfordian. In his description, he notes that the teeth of this specimen are similar to that of ''Ischyrodon'', but finally considered the two taxa as distinct in 1856. In 1883, Russian paleontologist Walerian Kiprijanoff suggested that ''Ischyrodon'' was closely related to ''
Pliosaurus ''Pliosaurus'' (meaning 'more lizard') is an extinct genus of thalassophonean pliosaurid known from the Late Jurassic (Kimmeridgian and Tithonian stages) of Europe and South America. This genus has contained many species in the past but recent ...
'', and consequently classified the latter two, along with ''
Polyptychodon ''Polyptychodon'' (meaning 'many-folded tooth') is a genus of pliosaurid found in Middle-Late Cretaceous marine deposits in southern England, France and Argentina. It has been considered a ''nomen dubium'' in a 2016 review. History of discover ...
'' and ''Thaumatosaurus'', within a new group of
sauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
ns that he named Thaumatosauria. Two years later,
Friedrich August von Quenstedt Friedrich August von Quenstedt (10 July 1809 – 21 December 1889) was a German geologist and palaeontologist. Life Von Quenstedt was born at Eisleben in Saxony, and educated at the Humboldt University of Berlin. After a period as assistant i ...
classified ''Ischyrodon'' among the
Plesiosauria The Plesiosauria or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period, possibly in the Rhaetian stage, about 203 million year ...
and highlighted its dental similarities with those of ''Pliosaurus'' and ''Polyptychodon''. While
Karl Alfred von Zittel Karl Alfred Ritter von Zittel (25 September 1839 – 5 January 1904) was a German palaeontologist best known for his ''Handbuch der Palaeontologie'' (1876–1880). Biography Karl Alfred von Zittel was born in Bahlingen in the Grand Duchy ...
considers ''Ischyrodon'' to be similar to ''Pliosaurus'', British paleontologist
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...
considered it a
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of this latter in 1889. He noted that the tooth had all the features present in teeth from
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
-aged England pliosaurs, and because of its large size he provisionally assigned it to the imposing species ''Pliosaurus macromerus''. Lydekker's opinion was thus followed by some studies published during the 20th century. In 1960, Lambert Beverly Tarlo suggested that the tooth of ''Ischyrodon'' seemed to have similar traits to that of ''
Liopleurodon ''Liopleurodon'' (; meaning 'smooth-sided teeth') is an extinct genus of carnivorous pliosaurid pliosaurs that lived from the Callovian stage of the Middle Jurassic to the Kimmeridgian stage of the Late Jurassic period (c. 166 to 155 mya). T ...
'' and was provisionally considered as a synonym of it. However, the author notes that his affection is not entirely justified based on the figure present in von Meyer's 1856 description. Tarlo's view was followed in many subsequent studies and works, noting that this tooth would be a tangible proof of the presence of ''Liopleurodon'' in Switzerland. The rise of studies describing
pliosaurid Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
teeth from the
thalassophonea Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
n subgroup prompted Czech paleontologist Daniel Madzia and colleagues to revise this synonymy in 2022. The authors noted that while the tooth most likely came from a ''Liopleurodon'' or a similar animal, they find that there is too little information available to make a confident assignment, leaving then ''Ischyrodon'' as a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
''.


Description

''Ischyrodon'' is a plesiosaurian of the "pliosauromorph" morphotype, meaning it had a large head and a short neck, unlike the "plesiosauromorphs", in which these anatomical proportions are reversed. Like all other plesiosaurs, ''Ischyrodon'' had a short tail, a barrel-shaped body, and all of its limbs modified into large flippers. The large size of the holotype tooth crown of ''Ischyrodon'' suggests that it came from a particularly large pliosaurid, although no size estimate has been given. The morphology of the latter nevertheless suggests that it originates from the anterior half of the right
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
or from the left side of the
dentary bone In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
. The dental crown also has apicobasal ridges, whose function was to improve the gripping and perforation of prey when biting. However, since ''Ischyrodon'' is recognized as a ''nomen dubium'', it possesses traits that are indistinguishable from other thalassophonean pliosaurids.


Paleoecology and dating

Although correctly noted as dating from Middle Jurassic since 1838, there was a uncertainty as to which stratigraphic stage the tooth exactly came from. In 1856, von Meyer noted that the specimen was of "Broadfordian" age. In 1889, Lydekker noted that based on its anatomy, the tooth was Kimmeridgian-aged, but this was later disputed by Tarlo in 1960, noting that its round anatomy in cross-section and associated
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
fossils suggested that it was possibly Oxfordian. However, more recent studies suggest that fossils found in the vicinity of the holotype tooth, as well as rocks in the iron mine from which it was probably extracted, are more likely to be
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 165.3 ± 1.1 Ma (million years ago) and 161.5 ± 1.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
-aged. The sedimentary
facies In geology, a facies ( , ; same pronunciation and spelling in the plural) is a body of rock with distinctive characteristics. The characteristics can be any observable attribute of rocks (such as their overall appearance, composition, or con ...
still present on the holotype tooth proves that it was preserved in a red
iron Iron is a chemical element; it has symbol Fe () and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, forming much of Earth's o ...
oolite Oolite or oölite () is a sedimentary rock formed from ooids, spherical grains composed of concentric layers. Strictly, oolites consist of ooids of diameter 0.25–2 millimetres; rocks composed of ooids larger than 2 mm are called pis ...
, which, within the
Fricktal The Fricktal ("Frick Valley") is a region on Northwestern Switzerland, comprising the Laufenburg and Rheinfelden districts of the Swiss canton of Aargau. The region was known as ''Frickgau'' in the medieval period, ultimately from a Late Lati ...
region, are rocks that are exclusively dated to the
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 165.3 ± 1.1 Ma (million years ago) and 161.5 ± 1.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
. However, erroneous references from miners indicated that the specimen would came from the
Hauptrogenstein Formation The Hauptrogenstein Formation is a Formation (geology), geologic formation in Germany and Switzerland. It preserves fossils that date back to the Middle Jurassic Period (geology), period (Bajocian-Bathonian). Paleofauna List as per Miedema ''e ...
, suggesting a
Bajocian In the geologic timescale, the Bajocian is an age and stage in the Middle Jurassic. It lasted from approximately 170.9 ±0.8 Ma to around 168.2 ±1.2 Ma (million years ago). The Bajocian Age succeeds the Aalenian Age and precedes the Bathonia ...
to
Bathonian In the geologic timescale the Bathonian is an age (geology), age and stage (stratigraphy), stage of the Middle Jurassic. It lasted from approximately 168.2 ±1.2 annum, Ma to around 165.3 ±1.1 Ma (million years ago). The Bathonian Age succeeds ...
age. Although plesiosaurs and
ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s have been found in this formation, this latter contains rocks that are generally gray in color, thus not fitting with the facies of the ''Ischyrodon'' tooth. Two ore beds located around Herznach and Wölflinswil preserve abundant and well-preserved
ammonite Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
fossils, corresponding to the Ifenthal Formation and probably also dating from the Callovian. Younger sedimentary rocks dating from the Oxfordian may also contain
ooid Ooids (, ) are small (commonly ≤2 mm in diameter), spheroidal, "coated" (layered) sedimentary grains, usually composed of calcium carbonate, but sometimes made up of iron- or phosphate-based minerals. Ooids usually form on the sea floo ...
s and can be quite rich in iron. However, these tend to be more grayish to greenish in color.


See also

*
List of plesiosaur genera This list of plesiosaurs is a comprehensive listing of all genera that have ever been included in the order Plesiosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered inv ...
*
Timeline of plesiosaur research This timeline of plesiosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, Taxonomy (biology), taxonomic revisions, and cultural portrayals of plesiosaurs, an order of marine reptiles ...


References

{{Taxonbar, from=Q113549840 Pliosauridae Middle Jurassic plesiosaurs of Europe Fossil taxa described in 1838 Sauropterygian genera