Polyodon Tuberculata
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''Polyodon tuberculata'' is an extinct species of
acipenseriform Acipenseriformes is an order of basal ray-finned fishes that includes living and fossil sturgeons and paddlefishes (Acipenseroidei), as well as the extinct families Chondrosteidae and Peipiaosteidae. They are the second earliest divergin ...
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
. It has only been found in sites in
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
dating back to less 1 million years after the Cretaceous-Paleocene Extinction. It is larger than the
American paddlefish The American paddlefish (''Polyodon spathula''), also known as a Mississippi paddlefish, spoon-billed cat, or spoonbill, is a species of ray-finned fish. It is the last extant taxon, living species of paddlefish (Polyodontidae). This family is ...
(''Polyodon spathula''), the only extant species of its genus, with the largest specimen being from a 2.6 meter long individual. The most notable feature of P. tuberculata is the presence of
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projectio ...
s that would have been weakly attached to the "paddle" and
skull roof The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In com ...
. Just like the modern species, ''P. tuberculata'' was a
filter-feeder Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a specia ...
in freshwater environments.


History and naming

The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
(
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 a ...
130629) is made up of a mostly-complete
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 ...
skull along with an associated upper caudal lobe and was first referenced by Laurie J. Bryant in 1989 as "Polyodontidae, undescribed". This specimen was collected in 1976 from the Farrand Channel of the upper
Tullock Formation The Tullock Member is a geologic member in Montana and Wyoming, forming the lowermost part of the Fort Union Formation. It is also known less commonly as the Tullock Formation, forming part of the Fort Union Group.. It preserves fossils dating b ...
located in
Garfield County, Montana Garfield County is a county located in the U.S. state of Montana. As of the 2020 census, the population was 1,173. Its county seat is Jordan. Garfield County is noteworthy as the site of the discovery and excavation of four of the world's doz ...
by Harley Garbani. At the time, the material was being studied by W. Bemis at the
University of Massachusetts The University of Massachusetts is the Public university, public university system of the Massachusetts, Commonwealth of Massachusetts. The university system includes six campuses (Amherst, Boston, Dartmouth, University of Massachusetts Lowell ...
. In this first mention of the material, the specimen known as LACM 126130 was considered as similar to the holotype, though it was later believed to be a specimen of '' Paleopsephurus wilsoni'' by multiple authors. The holotype would later be described in detail by Grande & Bemis (2010) from the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educationa ...
and University of Massachusetts respectively. More specimens from the
Fort Union Formation The Fort Union Formation is a geologic unit containing sandstones, shales, and coal beds in Wyoming, Montana, and parts of adjacent states. In the Powder River Basin, it contains important economic deposits of coal, uranium, and coalbed methane. ...
were later be described by Murray
et al. References Notes References Further reading * * External links * {{Latin phrases E ...
in 2020 based on three specimens collected during fieldwork in 2014 and 2015. These specimens are also made up of skull material, though one (UWBM 109828) also contains the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
. The species name "'' tuberculata''" refers to the
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projectio ...
s present on the holotype specimen's
skull roof The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In com ...
.


Description

The overall anatomy of ''Polyodon tuberculata'' is very similar to what is seen in the modern species, '' P. spathula''; the fossil fish had a long skull that was mostly made up of an extended "paddle" formed by a number of median rostral bones along with a network of
stellate Stellate, meaning star-shaped, may be used to describe: Biology * Stellate cell (disambiguation) * Stellate ganglion * Stellate reticulum * Stellate veins * Stellate trichomes (hairs) Other * Stellate wounds from lacerations or incisions * S ...
bones; though there are fewer stellate bones than what is seen in ''P. spathula''. A complete rostrum is not known from the species though it was most likely similar to what is seen in the modern species, with it not thinning anteriorly. The skull roof of P. tuberculata is also similar to its extant relative though it does possess a series of tubercles and crests on the post- temporal, frontals, and parietals. Due to these tubercles not being present on specimens besides the holotype, it has been suggested that they would have been weakly attached to the skull roof and "paddle". The sub-
opercle The operculum is a series of bones found in bony fish and chimaeras that serves as a facial support structure and a protective covering for the gills; it is also used for respiration and feeding. Anatomy The opercular series contains four bon ...
is the only bone of the opercular series preserved in specimens and is more
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
than in ''P. spathula''. Due to the back of the sub-opercle having a
scalloped Scallop () is a common name that encompasses various species of marine bivalve molluscs in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related familie ...
shape, it has been suggested that there would have been splint-like projections in this area. The
gill arch Branchial arches or gill arches are a series of paired bony/ cartilaginous "loops" behind the throat ( pharyngeal cavity) of fish, which support the fish gills. As chordates, all vertebrate embryos develop pharyngeal arches, though the event ...
es of the fish are mostly known from articulated
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of th ...
s that were overall thin. Based on the size of specimens, ''P. tuberculata'' was larger than its modern relative with the largest specimen most likely having a body length of . Not much of the
postcrania The postcranium ("behind the cranium"; plural: postcrania) or postcranial skeleton in zoology and vertebrate paleontology is the skeleton apart from the skull. The postcranium encompasses the axial skeleton, which includes the entirety of the verte ...
of the fish is known outside of the pectoral area and a small piece of the
caudal fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only ...
. Unlike the living species, the first ray of the pectoral fin is the same thickness as the rest of the rays though it is still the longest ray. Based on the small amount of the caudal fin preserved, the shape would have been very similar to what is seen in ''P. spathula''. Like other acipenseriforms, rhombic scales would have been present on the upper lobe of the caudal fin.


Paleobiology

Based on the presence of elongated
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of th ...
s and toothless jaws on specimens of ''P. tuberculata'', it is suggested that these fish would have been
filter feeder Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a s ...
s similar to their extant relative. Both formations that bear these specimens are representative of freshwater ecosystems with the fish being found in deposits less than 1 million years after the Cretaceous-Paleocene Extinction. The climate of Montana during this time would have been seasonally wet with the flora found at the Fort Union Formation suggesting a
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
environment.
Angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
s made up almost all of the known flora with "''
Carya Hickory is a common name for trees composing the genus ''Carya'', which includes 19 species accepted by ''Plants of the World Online''. Seven species are native to southeast Asia in China, Indochina, and northeastern India (Assam), and twelve ...
"'' and ''
Macginitiea ''Macginitiea'' is an extinct genus in the family Platanaceae ranging from the Paleogene, Late Paleocene to Late Eocene of North America, known from the John Day Fossil Beds National Monument#Geology and paleontology, Clarno Formation of central ...
'' being the most common plant
macrofossil Macrofossils, also known as megafossils, are the preserved remnants of organic beings and their activities that are large enough to be visible without a microscope. The term ''macrofossil'' stands in opposition to the term microfossil. Microfoss ...
s found. Based on various aspects of the geology of the formation, the
delta Delta commonly refers to: * Delta (letter) (Δ or δ), the fourth letter of the Greek alphabet * D (NATO phonetic alphabet: "Delta"), the fourth letter in the Latin alphabet * River delta, at a river mouth * Delta Air Lines, a major US carrier ...
of the
Catatumbo River The Catatumbo River () is a river rising in northern Colombia, flowing into Lake Maracaibo in Venezuela. The Catatumbo River is approximately long. It forms a part of the international boundary between the two countries. The river flows through ...
and
Lake Maracaibo Lake Maracaibo () is located in northwestern Venezuela, between the states of Zulia, Trujillo, and Mérida. While Maracaibo is commonly referred to as a lake, its current hydrological characteristics may better classify it as estuary and/or ...
have been suggested to be comparable to the bodies of water preserved at the formation. These bodies of water and those found at the Tullock Formation would have been a part of
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
s made up of
meander A meander is one of a series of regular sinuous curves in the Channel (geography), channel of a river or other watercourse. It is produced as a watercourse erosion, erodes the sediments of an outer, concave bank (cut bank, cut bank or river cl ...
ing channels, similar to the underlying
Hell Creek Formation The Hell Creek Formation is an intensively studied division of mostly Upper Cretaceous and some lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana. The Formation (stratigraphy), formation s ...
.
Backswamp In geology, a backswamp is a type of depositional environment commonly found in a floodplain. It is where deposits of fine silts and clays settle after a flood A flood is an overflow of water (list of non-water floods, or rarely other flui ...
s are also preserved in the form of
lignite Lignite (derived from Latin ''lignum'' meaning 'wood'), often referred to as brown coal, is a soft, brown, combustible sedimentary rock formed from naturally compressed peat. It has a carbon content around 25–35% and is considered the lowest ...
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 ...
at the Tullock Formation.


References

{{taxon bar Polyodontidae Prehistoric ray-finned fish