Toyotamaphimeia Skeletal
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''Toyotamaphimeia machikanensis'' (
Toyotama-hime or Luxuriant-Jewel-Princess is a goddess in Japanese mythology in the episode of the "Luck of the Sea and the Luck of the Mountain" in the ''Kojiki'' as well as '' Nihon Shoki''. She is the daughter of the sea deity, Watatsumi. Toyotama marries t ...
from Mountain Machikane ( :ja:待兼山)) is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
gavialid Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii''), both occurring in Asia. Man ...
crocodylia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
n which lived in
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
during the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
. A specimen recovered in 1964 at
Osaka University , abbreviated as , is a public research university located in Osaka Prefecture, Japan. It is one of Japan's former Imperial Universities and a Designated National University listed as a "Top Type" university in the Top Global University Project. ...
during the construction of a new science building has been dated to around 430–380 thousand years old based on the stratum in which it was found. Unassigned species from same genus is also known from
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the nort ...
. ''T. machikanensis'' was a fairly large
crocodylian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living re ...
with a 1 m (3.3 ft) skull and a total length up to 7.7 m (25 ft). It was originally described as a member of the genus ''
Tomistoma ''Tomistoma'' is a genus of gavialid crocodilians. They are noted for their long narrow snouts used to catch fish, similar to the gharial. ''Tomistoma'' contains one extant (living) member, the false gharial (''Tomistoma schlegelii''), as we ...
''.


History and naming

The first bones belonging to ''Toyotamaphimeia'' were discovered on May 3rd 1964 during the construction of a new school building on the grounds of Ôsaka University. A field survey was conducted shortly afterwards, confirming the presence of more fossils, however not yet identifying their crocodilian nature. Following the survey several digs were organized starting on the 9th of June 1964. The skull was found on the 17th of September during the 2nd dig. A third excavation was held in December which yielded more material of ''Toyotamaphimeia'' as well as fossil shellfish, insects and plant remains. Finally a 4th excavation took place in January of 1965. Following analysis of the fossils, the material was assigned to the genus ''Tomistoma'' and named ''Tomistoma machikanense''. In 1983, 18 years after the initial discovery, the skull was redescribed and deemed different enough from ''Tomistoma'' to erect a new genus, ''Toyotamaphimeia'' creating the comb. nov. ''Toyotamaphimeia machikanensis''. In turn Aoki also changed the species name from ''machikanense'' to ''machikanensis'', as the new genus name was feminine. The generic name derives from Toyotama-hime, a goddess of Japanese mythology with the ability to change her appearance to that of a crocodile. The species epithet means "from Mountain Machikane".


Description

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 ...
of ''Toyotamaphimeia'' is a nearly complete skeleton consisting of a skull, an entire cervical and dorsal series of vertebrae, various ribs, 33
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...
as well as almost half the bones of the limbs, hip region and pectoral girdle. Most of the tail past the 3rd cervical vertebra is missing, making it difficult to determine the exact length of the animal. ''Toyotamaphimeias skull is triangular in shape and longirostrine. It's fairly large, 1.025 meters from the tip of the premaxillary to the posterior end of the parietal. Most of that length is taken up by the maxilla and the nasal bones penetrate the premaxilla dorsally, extending deep into the premaxilla to the level of the 3rd maxillary alveoli, but not coming in contact with the nares. The skulltable of the holotype is crushed and damaged just before the orbits. The dentaries are broken off at the anterior end and each preserves 10 alveoli. The absence of any grooves or confluence of alveoli suggests that the specimen is mature, which is consistent with its great size.


Paleobiology

The holotype specimen (MOUF00001) preserves a series of pathologies described by Katsura in 2004. The mandible is broken off at the tip, the tibia and fibula have been fractured and healed and some of the osteoderms present preserve healing bite marks. The fact that these injuries healed is evidence that the animal survived for a while after being injured and Katsura suggests that they may have been the result of intraspecific fights, furthermore hypothesizing that this could mean the Osaka University specimen may have been a male. Although the holotype of ''Toyotamaphimeia'' is the first substantial and best preserved evidence of crocodilians in Japan, there are other remains found across the islands. The northernmost finds were made in the Iwate Prefecture (northern
Honshu , historically called , is the largest and most populous island of Japan. It is located south of Hokkaidō across the Tsugaru Strait, north of Shikoku across the Inland Sea, and northeast of Kyūshū across the Kanmon Straits. The island separ ...
) while their range extends south to Nagasaki Prefacture (
Kyushu Island is the third-largest island of Japan's five main islands and the most southerly of the four largest islands ( i.e. excluding Okinawa). In the past, it has been known as , and . The historical regional name referred to Kyushu and its surroun ...
). At this latitude ''Toyotamaphimeia'' would have existed at the thermal limit of crocodilians. The Ibaraki Formation, where the remains of ''Toyotamaphimeia'' have been found, is part of the Osaka Group, which consists of lacustrine and fluvial deposits of the Pliocene to Pleistocene. Specifically, the fossils belong to the Kasuri Tuff, which dates to the
Chibanian The Chibanian, widely known by its previous designation of Middle Pleistocene, is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocene Epoch withi ...
age of the Pleistocene. Molluscs, pollen and plant fossils (species of lotus and
water caltrop The water caltrop is any of three extant species of the genus ''Trapa'': ''Trapa natans'', ''Trapa bicornis'' and the endangered ''Trapa rossica''. It is also known as buffalo nut, bat nut, devil pod, ling gok (Chinese: 菱角), ling nut, lin kok ...
found in the Kasuri Tuff suggest a moderate climate. ''Toyotamaphimeia'' would have most likely coexisted in this area alongside '' Stegodon orientalis'', ''
Cervus kazusensis ''Cervus'' is a genus of deer that primarily are native to Eurasia, although one species occurs in northern Africa and another in North America. In addition to the species presently placed in this genus, it has included a whole range of other sp ...
'', ''
Panthera youngi ''Panthera youngi'' is a fossil cat species that was described in 1934; fossil remains of this cat were excavated in a ''Sinanthropus'' formation in Choukoutien, northeastern China. Upper and lower jaws excavated in Japan's Yamaguchi Prefecture ...
'', ''
Bubalus teihardi ''Bubalus'' is a genus of Asiatic bovines that was proposed by Charles Hamilton Smith in 1827. ''Bubalus'' and ''Syncerus'' form the subtribe Bubalina, the true buffaloes. The International Code of Zoological Nomenclature and classification of d ...
'' and ''
Stephanorhinus ''Stephanorhinus'' is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Pliocene to Late Pleistocene. Species of ''Stephanorhinus'' were the predominant and often only species of rhinoceros in much ...
''. The pollen found in the region suggests a vegetation consisting of
alder Alders are trees comprising the genus ''Alnus'' in the birch family Betulaceae. The genus comprises about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few sp ...
s,
beeches Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...
,
pines A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
and ''
Cryptomeria ''Cryptomeria'' (literally "hidden parts") is a monotypic genus of conifer in the cypress family Cupressaceae, formerly belonging to the family Taxodiaceae. It includes only one species, ''Cryptomeria japonica'' ( syn. ''Cupressus japonica'' L ...
'' (Japanese redwood).


Phylogeny

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 ...
based morphological studies comparing skeletal features that shows ''Toyotamaphimeia'' as a member of
Tomistominae Tomistominae is a subfamily of crocodylians that includes one living species, the false gharial. Many more extinct species are known, extending the range of the subfamily back to the Eocene epoch. In contrast to the false gharial, which is a fre ...
, related to the
false gharial The false gharial (''Tomistoma schlegelii''), also known by the names Malayan gharial, Sunda gharial and tomistoma is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Vu ...
: Based on morphological studies of
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
taxa In biology, a taxon (back-formation from ''taxonomy''; plural 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 nam ...
, the tomistomines (including the living
false gharial The false gharial (''Tomistoma schlegelii''), also known by the names Malayan gharial, Sunda gharial and tomistoma is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Vu ...
) were long thought to be classified as
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s and not closely related to
gavialoids Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial ''Gavialis gangeticus'' and the false gharial ''Tomistoma schlegelii'' are ...
. However, recent molecular studies using
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
have consistently indicated that the
false gharial The false gharial (''Tomistoma schlegelii''), also known by the names Malayan gharial, Sunda gharial and tomistoma is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Vu ...
(''Tomistoma'') (and by inference other related extinct forms in
Tomistominae Tomistominae is a subfamily of crocodylians that includes one living species, the false gharial. Many more extinct species are known, extending the range of the subfamily back to the Eocene epoch. In contrast to the false gharial, which is a fre ...
) actually belong to
Gavialoidea Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial ''Gavialis gangeticus'' and the false gharial ''Tomistoma schlegelii'' are ...
(and
Gavialidae Gavialidae is a family (biology), family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two extant taxon, living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii ...
). 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 ...
from a 2018
tip dating Tip dating is a technique used in molecular dating that allows the inference of time-calibrated phylogenetic trees. Its defining feature is that it uses the ages of the samples to provide time information for the analysis, in contrast with traditio ...
study by Lee & Yates simultaneously using morphological, molecular (
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
), and
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostrati ...
(
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 preserved ...
age) data that shows ''Toyotamaphimeia'' as a
gavialid Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii''), both occurring in Asia. Man ...
, related to both the
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
and the
false gharial The false gharial (''Tomistoma schlegelii''), also known by the names Malayan gharial, Sunda gharial and tomistoma is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Vu ...
:


References


External links

* * {{Taxonbar, from=Q6130295, from2=Q15041708 Gavialidae Fossil taxa described in 1983 Fossils of Japan Prehistoric pseudosuchian genera Pleistocene crocodylomorphs Pleistocene reptiles of Asia Pleistocene genus first appearances