Tinirau (genus)
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''Tinirau'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
sarcopterygian Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...
fish from the
Middle Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wher ...
of
Nevada Nevada ( ; ) is a U.S. state, state in the Western United States, Western region of the United States. It is bordered by Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. N ...
. Although it spent its entire life in the ocean, ''Tinirau'' is a stem tetrapod close to the ancestry of land-living vertebrates in the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
Tetrapoda. Relative to more well-known stem tetrapods, ''Tinirau'' is more closely related to Tetrapoda than is ''
Eusthenopteron ''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
'', but farther from Tetrapoda than is ''
Panderichthys ''Panderichthys'' is a genus of extinct sarcopterygian (lobe-finned fish) from the late Devonian period, about 380 Mya. ''Panderichthys'', which was recovered from Frasnian (early Late Devonian) deposits in Latvia, is represented by two species. ...
''. The type and only species of ''Tinirau'' is ''T. clackae'', named in 2012.


Description

''Tinirau'' was a fairly large, predatory fish about a meter long and with a deep, compact body. The head was large, with large terminal mout and numerous teeth. The tail was
heterocercal Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
, but the remaining fins with the exception of the
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
s was situated behind the middle of the body similar to the situation seen in pikes, giving the animal a large tail surface suitable for great bursts of speed. It shares many advanced features with later tetrapodomorphs in the pelvic limb bones and
glenoid The glenoid fossa of the scapula or the glenoid cavity is a bone part of the shoulder. The word ''glenoid'' is pronounced or (both are common) and is from el, gléne, "socket", reflecting the shoulder joint's ball-and-socket form. It is a sha ...
s (shoulder sockets). By the time ''Tinirau'' appeared, many stem tetrapods had already developed the three major hindlimb bones of tetrapods: the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
,
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, and
fibula The fibula 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. Its upper extremity is ...
. While derived stem tetrapods like ''Panderichthys'' had hindlimb configurations very similar to the first land-living tetrapods, some early forms such as ''
Eusthenopteron ''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
'' possessed a prominent postaxial process of the
fibula The fibula 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. Its upper extremity is ...
hanging over the fibulare bone below it. ''Tinirau'' is the earliest known stem tetrapod to have a significantly reduced postaxial process, and a fibula more like those of later tetrapods. Like those of the Late Devonian ''
Panderichthys ''Panderichthys'' is a genus of extinct sarcopterygian (lobe-finned fish) from the late Devonian period, about 380 Mya. ''Panderichthys'', which was recovered from Frasnian (early Late Devonian) deposits in Latvia, is represented by two species. ...
'' and ''
Ichthyostega ''Ichthyostega'' (from el, ἰχθῦς , 'fish' and el, στέγη , 'roof') is an extinct genus of limbed tetrapodomorphs from the Late Devonian of Greenland. It was among the earliest four-limbed vertebrates in the fossil record, and was on ...
'', the glenoid of ''Tinirau'' is elongated along the anteroposterior (forward-backward) axis of the body. The lengthening of the glenoid corresponds with a flattening of the proximal end of the humerus, a feature common in the forelimbs of more advanced stem tetrapods. Although the pectoral limb bones and girdle were not strong enough to support the weight of ''Tinirau'' out of water, glenoid lengthening and other changes to the proximal forelimb were among the first steps in the transformation from pectoral fin to forelimb.


History

Remains of ''Tinirau'' were first discovered by paleontologist
Joseph T. Gregory Dr. Joseph Tracy Gregory (July 28, 1914 – November 18, 2007) was an American paleontology, paleontologist and professor. Joseph Tracy Gregory was born in Eureka, California, the only child of Frank Gregory, a civil engineer, and Edith Tracy, ...
in 1970, who then worked for the
University of California Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant univ ...
. Gregory and his field team found these fossils near the
Simpson Park Mountains The Simpson Park Mountains, also known as the Simpson Park Range, are located in Lander and Eureka counties, in central Nevada in the western United States. The mountains reach an elevation of at Fagin Mountain, and cover an area of approximat ...
in
Eureka County, Nevada Eureka County is a county in the U.S. state of Nevada. As of the 2020 census, the population was 1,855, making it the second-least populous county in Nevada. Its county seat is Eureka. Eureka County is part of the Elko Micropolitan Statistic ...
. They came from a deposit called the Red Hill I beds, which date to the upper
Givetian The Givetian is one of two faunal stages in the Middle Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Eifelian Stage and followed by the Frasnian Stage. It is named after the town of Givet in Fra ...
stage of the Middle Devonian. The Red Hill I beds include a series of
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
and
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
, likely deposited in an
outer continental shelf The Outer Continental Shelf (OCS) is a feature of the geography of the United States. The OCS is the part of the internationally recognized continental shelf of the United States which does not fall under the jurisdictions of the individual U. ...
marine environment. Six fossils are known, all preserving bones of the skull. Two fossils preserve postcranial bones in articulation. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
specimen,
UCMP The University of California Museum of Paleontology (UCMP) is a paleontology museum located on the campus of the University of California, Berkeley. The museum is within the Valley Life Sciences Building (VLSB), designed by George W. Kelham and ...
118605, is a mostly complete skeleton. ''Tinirau clackae'' was named by paleontologist Brian Swartz in 2012 after
Tinirau In Polynesian mythology, stories about Tinirau are found throughout the islands of Polynesia. He is a guardian of fish. Many themes recur in the various versions. Often he travels to another land in search of his wife, or his wife travels to anot ...
, who in Polynesian legend is the guardian of ocean life, and whose form is half human and half fish. The connection alludes to the features in ''Tinirau'' that are transitional between fish, representing life in the water, and tetrapods, representing life on land. The specific name honors
Jennifer A. Clack Jennifer Alice Clack, (''née'' Agnew; 3 November 1947 – 26 March 2020) was an English palaeontologist and evolutionary biologist. She specialised in the early evolution of tetrapods, specifically studying the "fish to tetrapod" transition: ...
, an English paleontologist who has made many contributions to the study of stem tetrapods.


Phylogeny

In
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
terms, ''Tinirau'' is positioned more crownward, or closer to Tetrapoda, than other stem tetrapods like
megalichthyiform Megalichthyiformes is an extinct clade of basal tetrapodomorphs which first appeared during the Devonian period. It was named in 2010 by Michael I. Coates and Matt Friedman, who defined it as a stem-based taxon containing all tetrapodomorphs clo ...
s and
tristichopterid Tristichopterids (Tristichopteridae) were a diverse and successful group of tetrapodomorph fishes living throughout the Middle and Late Devonian. They first appeared in the Eifelian stage of the Middle Devonian. Within the group sizes ranged from ...
s. In its first description, the phylogenetic analysis of Swartz placed ''Tinirau'' as a branch directly connected to the stem of Tetrapoda. In other words, it is a close relative of the lineage leading to land vertebrates (a prehistoric taxon can rarely be identified as an ancestor of another taxon, but is instead considered a side branch in a larger lineage). ''Eusthenopteron'', traditionally considered a direct ancestor of tetrapods, or a close relative of such an ancestor, was placed deep within the family
Tristichopteridae Tristichopterids (Tristichopteridae) were a diverse and successful group of tetrapodomorph fishes living throughout the Middle and Late Devonian. They first appeared in the Eifelian stage of the Middle Devonian. Within the group sizes ranged from ...
, away from the tetrapod stem. Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
modified from Swartz (2012) showing the placement of ''Tinirau'': On the other hand, Parfitt ''et al.'' (2014) considered it more likely that ''Tinirau'' was a member of the family Tristichopteridae.


References

{{Taxonbar, from1=Q290682, from2=Q20719461 Eotetrapodiforms Devonian animals Devonian fish of North America