Rhizodus
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''Rhizodus'' ("root tooth") is an extinct genus of basal, finned
tetrapodomorph Tetrapodomorpha (also known as Choanata) is a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced for ...
s (the group of
sarcopterygian Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
s that contains modern
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s and their extinct relatives). It belonged to
Rhizodontida Rhizodontida is an extinct group of predatory tetrapodomorphs known from many areas of the world from the Givetian through to the Pennsylvanian - the earliest known species is about 377 million years ago (Mya), the latest around 310 Mya. Rhizod ...
, one of the earliest-diverging tetrapodomorph
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s. Two valid species have been described, both of which lived during the
Early Carboniferous Early may refer to: Places in the United States * Early, Iowa, a city * Early, Texas, a city * Early Branch, a stream in Missouri * Early County, Georgia * Fort Early, Georgia, an early 19th century fort Music * Early B, stage name of Jamaican d ...
epoch. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''R. hibberti'' is known from the
Viséan The Visean, Viséan or Visian is an age in the ICS geologic timescale or a stage in the stratigraphic column. It is the second stage of the Mississippian, the lower subsystem of the Carboniferous. The Visean lasted from to Ma. It follows ...
stage of the United Kingdom, whereas the species ''R. serpukhovensis'' is from the
Serpukhovian The Serpukhovian is in the ICS geologic timescale the uppermost stage or youngest age of the Mississippian, the lower subsystem of the Carboniferous. The Serpukhovian age lasted from Ma to Ma. It is preceded by the Visean and is followed ...
of Russia. Some fossils referred to the genus ''Rhizodus'' have also been found in North America.


History

A partial lower jaw attributed to a rhizodontiform, PIN no. 2878/581, was discovered in 2000 in the Zaborie quarry of
Serpukhovsky District Serpukhovsky District () is an administrativeLaw #11/2013-OZ and municipalLaw #78/2005-OZ district (raion), one of the thirty-six in Moscow Oblast, Russia. It is located in the south of the oblast. The area of the district is . Its administrative ...
, Russia. The specimen consists of the middle part of the lower jaw, which includes the two coronoid fangs as well as marginal teeth. In 2022, it was formally described as the holotype of the new species ''R. serpukhovensis''. Dating from the
Serpukhovian The Serpukhovian is in the ICS geologic timescale the uppermost stage or youngest age of the Mississippian, the lower subsystem of the Carboniferous. The Serpukhovian age lasted from Ma to Ma. It is preceded by the Visean and is followed ...
, it was the youngest known specimen of ''Rhizodus'', and the first known from Russia. No near-complete adult skeleton of ''Rhizodus'' has been found, and the known material is limited to isolated fragments. Given explanations include
taphonomic bias Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov ...
due to large body size, but also weak
endochondral ossification Endochondral ossification is one of the two essential pathways by which bone tissue is produced during fetal development and bone healing, bone repair of the mammalian skeleton, skeletal system, the other pathway being intramembranous ossificatio ...
in rhizodontids, and a lack of strong
cranial sutures In anatomy, fibrous joints are joints connected by Fibrous connective tissue, fibrous tissue, consisting mainly of collagen. These are fixed joints where bones are united by a layer of white fibrous tissue of varying thickness. In the skull, the ...
between skull bones.


Description

The most notable characteristics of ''Rhizodus'', compared to other giant rhizodonts such as ''
Barameda ''Barameda'' (Indigenous Australian language: "fish trap") is a genus of rhizodont Sarcopterygii, lobe-finned fishes which lived during the Tournaisian stage near the start of the Carboniferous period in Australia; fossils of the genus have been ...
'', were the two fangs located near the front of its jaws, followed by other teeth scaling downwards in size. ''Rhizodus'' was a giant
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
that resided in freshwater lakes, river systems and large swamps, with ''R. hibberti'' measuring long and weighing . It fed on small to medium-sized amphibians, using its teeth to kill prey and rip it into digestible sizes, rather than swallowing prey whole like other, smaller-toothed
sarcopterygians Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
. Fossil skin imprints show that ''Rhizodus'' had large, plate-like scales, similar to those found on modern day
arapaima The arapaima, pirarucu, or paiche is any large species of bonytongue in the genus ''Arapaima'' native to the Amazon Basin, Amazon and Essequibo River, Essequibo basins of South America. ''Arapaima'' is the type genus of the subfamily Arapaiminae ...
. In ''Rhizodus'', the
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hi ...
s consisted of a
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
articulating with three radial bones. This is unlike modern sarcopterygians and other known fossils, where the femur articulates with two bones, the
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
and
fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...
, which is believed to be the basal condition. The femur and the three radial bones in ''Rhizodus'' are thought to originate respectively from the stylopod and zeugopod, although without homology between the individual radials in ''Rhizodus'' and those in other tetrapods. The pectoral fins comprised a
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
articulating with a
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
and
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
, making them closer to the tetrapod condition.


Taxonomy

''Rhizodus'' is part of the eponymous clade
Rhizodontida Rhizodontida is an extinct group of predatory tetrapodomorphs known from many areas of the world from the Givetian through to the Pennsylvanian - the earliest known species is about 377 million years ago (Mya), the latest around 310 Mya. Rhizod ...
, usually considered to be an early-diverging
tetrapodomorph Tetrapodomorpha (also known as Choanata) is a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced for ...
branch. However, the discovery of '' Hongyu'' in 2017, showing a mosaic of rhizodont-like and tetrapod-like characters, has questioned this placement. The phylogeny below comes from a study performed by Clement et al. in 2021, which recovered Rhizodontida as basal tetrapodomorphs positioned crownwards of ''
Kenichthys ''Kenichthys'' is a genus of sarcopterygian fish from the Devonian period, and a member of the clade Tetrapodomorpha. The only known species of the genus is ''Kenichthys campbelli'' (named for the Australian palaeontologist Ken Campbell), the f ...
'' and ''
Tungsenia ''Tungsenia'' is an extinct genus of basal tetrapodomorph bony fish known from the ~409 mya (Early Devonian) of northeastern Yunnan Province, China China, officially the People's Republic of China (PRC), is a country in East Asia. With ...
''. While the genera ''Rhizodus'', ''Strepsodus'' and '' Archichthys'' are considered well-established, distinguishing characters focus on the symphysial tusks, leaving identification of other material more uncertain due to the fragmentary nature of remains. A 2020 study by Johanson et al. suggested dividing rhizodont
humeri The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of ...
into four morphogroups. Morphogroup III specimens, characterized by a large preaxial process extending dorso-ventrally, were identified with ''Rhizodus'', tentatively extending the range of the genus to the
Tournaisian The Tournaisian is in the ICS geologic timescale the lowest stage or oldest age of the Mississippian, the oldest subsystem of the Carboniferous. The Tournaisian age lasted from Ma to Ma. It is preceded by the Famennian (the uppermost st ...
.


Diet

The diet of ''Rhizodus'' included medium-sized fish and tetrapods. It has been proposed that ''Rhizodus'' lunged at terrestrial, shorebound prey, just like a modern-day crocodile.


References


Works cited

* * * * * {{Taxonbar, from=Q7320749 Rhizodontida Prehistoric lobe-finned fish genera Viséan genera Serpukhovian life Mississippian sarcopterygians of Europe Mississippian sarcopterygians of North America Fossils of Great Britain Fossils of Russia Fossil taxa described in 1840 Taxa named by Richard Owen