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''Proganochelys'' is an extinct, primitive stem-turtle that has been hypothesized to be the sister
taxon In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechani ...
to all other turtles creating a
monophyletic group A clade (), also known as a monophyletic group or natural group, is a group of organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemist ...
, the ''Casichelydia''. ''Proganochelys'' was named by
Georg Baur Georg Baur (1859–1898) was a German Vertebrate paleontology, vertebrate paleontologist and Lamarckism, Neo-Lamarckian who studied reptile, reptiles of the Galapagos Islands, particularly the Galápagos tortoise, Galápagos tortoises, in the 1890s. ...
in 1887 as the oldest turtle in existence at the time. The name ''Proganochelys'' comes from the
Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as of ...
word ''ganos'' meaning brightness, combined with prefix ''pro'', and
Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as of ...
base ''chelys'' meaning "turtle". ''Proganochelys'' is believed to have been around 1 meter in size and
herbivorous A herbivore is an animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All ...

herbivorous
in nature. ''Proganochelys'' was known as the most primitive stem-turtle for over a
century A century is a period of 100 years. Centuries are numbered names of numbers in English#Ordinal numbers, ordinally in English and many other languages. The word ''century'' comes from the Latin ''centum'', meaning ''one hundred''. ''Century'' is s ...

century
, until the novel discovery of ''
Odontochelys ''Odontochelys semitestacea'' (meaning "toothed turtle with a half-shell") is a Late Triassic relative of turtles. Before ''Pappochelys'' was discovered and ''Eunotosaurus'' was redescribed, ''Odontochelys'' was considered the oldest undisputed m ...

Odontochelys
'' in 2008. ''
Odontochelys ''Odontochelys semitestacea'' (meaning "toothed turtle with a half-shell") is a Late Triassic relative of turtles. Before ''Pappochelys'' was discovered and ''Eunotosaurus'' was redescribed, ''Odontochelys'' was considered the oldest undisputed m ...

Odontochelys
'' and ''Proganochelys'' share unique primitive features that are not found in ''Casichelydia'', such as teeth on the
pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medial ...

pterygoid
and
vomer The vomer () is one of the unpaired facial bones The facial skeleton comprises the ''facial bones'' that may attach to build a portion of the skull The skull is a bone A bone is a Stiffness, rigid tissue (anatomy), tissue that constitute ...

vomer
and a plate-like coracoid. ''Proganochelys'' is the oldest stem-turtle species with a complete shell discovered to date, known from fossils found in
Germany ) , image_map = , map_caption = , map_width = 250px , capital = Berlin Berlin (; ) is the Capital city, capital and List of cities in Germany by population, largest city of Germany by both area and population. Its 3,769,495 inh ...

Germany
,
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an autonomous territory An autonomous administrative division (also referred to as an autonomous area, entity, unit, region, subdivision, or territory) is a subnational administra ...

Greenland
, and
Thailand Thailand ( th, ประเทศไทย), historically known as Siam, () officially the Kingdom of Thailand, is a country in Southeast Asia. It is located at the centre of the Mainland Southeast Asia, Indochinese Peninsula, spanning , wi ...

Thailand
in strata from the late
Triassic The Triassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...

Triassic
, dating to approximately 210 million years ago. The location of these fossils suggest that ''Proganochelys'' was active throughout the continent of
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea Pangaea or Pangea () was a supercontinent In geology Geology (from the Ancient Greek γῆ, ''gē'' ("earth") and -λoγία, ''-logia'', ("s ...
. There are only two known species of ''Proganchelys'', with little information as a result of a small fossil record. All ''Proganochelys quentesti'' fossils were discovered in Germany, while ''Proganochelys ruchae'' fossils were found in Thailand. ''Psammochelys'', ''Stegochelys'', and ''Triassochelys'' are junior synonyms of ''Proganochelys''. ''Chelytherium'' von Meyer, 1863 has been considered a synonym of ''Proganochelys'' by some authors, but Joyce (2017) considers it a ''nomen dubium'' given the fragmentary nature of the syntype material. Joyce (2017) also considered North American genus ''
Chinlechelys ''Chinlechelys'' ( meaning ''Chinle turtle'') is an extinct genus of stem-turtle belonging to Testudinata. It lived in the Norian age of the Late Triassic and is the oldest turtle known from North America. Among turtles it is unique, mostly becau ...
'' to be a junior synonym of ''Proganochelys'', though the author maintains the type species of the former genus, ''C. tenertesta'', as a distinct species within the genus ''Proganochelys''.


Description and paleobiology

''Proganochelys'' was once considered to be the oldest known stem-turtle until the description of ''
Odontochelys ''Odontochelys semitestacea'' (meaning "toothed turtle with a half-shell") is a Late Triassic relative of turtles. Before ''Pappochelys'' was discovered and ''Eunotosaurus'' was redescribed, ''Odontochelys'' was considered the oldest undisputed m ...

Odontochelys
'', a slightly earlier genus that lived in the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the sta ...
stage of the
Triassic The Triassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...

Triassic
. In life it was about 3 feet long with a fully developed shell. Its overall appearance resembled modern turtles in many respects: it lacked teeth on the upper and lower jaw, likely had a beak and had the characteristic heavily armored Turtle shell, shell formed from bony plates and ribs which fused together into a solid cage around the internal organs. ''Proganochelys'' had a semi-beak like structure along with teeth fused to its
vomer The vomer () is one of the unpaired facial bones The facial skeleton comprises the ''facial bones'' that may attach to build a portion of the skull The skull is a bone A bone is a Stiffness, rigid tissue (anatomy), tissue that constitute ...

vomer
. The plates comprising the carapace and plastron were already in the modern form, although there were additional plates along the margins of the shell that would have served to protect the legs. Also unlike any modern species of turtle, its long tail had spikes and terminated in a club, its head could not be retracted under the shell and its neck may have been protected by small spines. While it had no teeth in its jaws, it did have small Denticle (tooth feature), denticles on the palate. The beak like structure suggests that the Triassic stem-turtles evolved from carnivorous stem-turtles to
herbivorous A herbivore is an animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All ...

herbivorous
as the loss of teeth and gain of the beak would benefit the crushing of plants in these stem-turtles.


Synapomorphies and autapomorphies

''Proganochelys'' possess a few chelonian synapomorphies including: a bony shell containing fused ribs, neural bones with fused thoracic segments, and a carapace and plastron that enclose the pelvic and shoulder girdle. ''Proganochelys'' was also known for its autapomorphy features which included a tail club and a tubercle on the basioccipital. The tail of ''Proganochelys'' was noticeably long and is hypothesized to have been used as a club for protection against predators. Although evolution of the shell has been clearly defined, the mechanisms behind the movement of the neck has been a subject of debate for ''Proganochelys''. It has been hypothesized that ''Proganochelys'' were able to retract their necks by tucking in their skull under the front of their shell when needed.


Shell

The broadened ribs on ''Proganochelys'' show "metaplastic ossification of the dermis". The enlarged ribs suggest that the endochondral rib ossifications were joined by a second ossification instead of having expanded ribs. The 220-million-year-old stem-turtle ''Odontochelys'' only has a partially formed shell. ''Odontochelys'' is believed to only possess the underside element of a shell known as a plastron. The 5-million-year difference that distinguish ''Odontochelys'' from ''Proganochelys'' tell us that the evolution of the shell occurred relatively quickly in time. ''Proganochelys'' possess both a carapace, the upper formation of the shell, and the plastron, the lower. The shell is believed to be used for protection an enhanced feature for survival. ''Proganochelys'' fits well into the order as a turtle, as the shell of ''Proganochelys'' is in agreement with the evolution of other stem-turtles.


Skull

The dermal roofing elements of ''Proganochelys'' include a large nasal cavity, nasal, a fully roofed skull, a flat squamosal, and an absent pineal foramen. Palatal characteristics include paired vomers, and a dorsal process containing premaxilla. An open interpterygoid vacuity along with a prominent elongated quadrate are notable basicranial elements. Overall, ''Pragonchelys'' is characterized by having few chelonian features and having a relatively generalized amniote skull. The skull of ''Proganochelys Quenstedti'' from Trossingen, West Germany, retains a number of well-known amniote features not found in any other turtle. For instance, the lacrimal bone, supratemporal bone, and lacrimal duct are notable structures that are kept. Furthermore, some traits that are present in modern turtles are not present in ''Proganochelys'' and therefore must have come after the evolution of the shell. For instance, jaw differentiation, the fusion of the vomer, and the loss of the lacrimal are clear examples of traits that evolved after the evolution of the shell in ''Proganochelys''.


Discovery

The earliest fossils of ''Proganochelys'' were discovered in
Germany ) , image_map = , map_caption = , map_width = 250px , capital = Berlin Berlin (; ) is the Capital city, capital and List of cities in Germany by population, largest city of Germany by both area and population. Its 3,769,495 inh ...

Germany
in the rural towns of Halberstadt, Tübingen, and Trossingen. The fossils were found in an elaborate formation of shales, sandstones, and some limestone piles, with the formation believed to be between 220 and 205 million years old. Consensus among Geologists placed the fossils in the middle of the Norian, around 210 million years ago, although this is largely an estimate. In addition to ''Proganochelys'', the rock formations in Germany have also given fossils for the stem-turtle ''Proterochersis''.


Paleoecology

The specific ecology of the Late
Triassic The Triassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...

Triassic
stem-turtles has been disputed and a major point of disagreement for many years among scientists.
Triassic The Triassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...

Triassic
stem-turtles, including ''Proganochelys'', appear to have been both aquatic and terrestrial. Shell proportions are believed to be correlated to the environment in which a turtle lives in, seen in modern turtles today. Using this concept, scientists were able to infer on the habitat in which ''Proganochelys'' may have lived in. A comparison between modern turtles and ''Proganochelys'' found that it was not likely that stem-turtles had differentiated into specialized ecologies such as open water swimmers or solely terrestrial animal, terrestrial turtles in the Late Triassic period. If this is the case, a freshwater habitat would be the most likely environment for ''Proganochelys'' to have lived in. On the other hand, it is noted that some believe ''Proganochelys'' were solely terrestrial. Shell bone histology of extant turtles revealed congruence with terrestrial turtles for the earliest basal turtles, including ''Proganochelys'', taxa in one study. The common ancestry of all living turtles is believed to be aquatic, while the earliest turtles are believed to have lived in a terrestrial environment.


Environment and forelimbs

Forelimbs are believed to be a physical feature that reflects the preferences and adaptations to a specific environment, indicating the environment a turtle would be most likely to reside in. Based on morphology (biology), morphological data, ''Proganochelys'' is believed to have lived in a semi-aquatic environment, though a 2021 study groups it with tortoises and other terrestrial taxa. Turtles possessing short hands are believed to be most likely terrestrial, while turtles with long limbs are more likely to be aquatic. The majority of all testudines are short-handed and terrestrial, while all cheloniods are long-handed and aquatic animal, aquatic.


Classification

''Proganochelys'' belongs to the group of tetrapods with a shell known as Testudinata and is the oldest primitive stem turtle. The group does not include ''Odontochelys''. The cladogram below follows an analysis by Jérémy Anquetin (2012).


Habitat

''Proganochelys'' is considered to have lived in the giant continent
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea Pangaea or Pangea () was a supercontinent In geology Geology (from the Ancient Greek γῆ, ''gē'' ("earth") and -λoγία, ''-logia'', ("s ...
during the
Triassic The Triassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...

Triassic
period. The fossil records show that ''Proganochelys'' might have lived anywhere in between Thailand and Germany. During the
Triassic The Triassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...

Triassic
period,
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea Pangaea or Pangea () was a supercontinent In geology Geology (from the Ancient Greek γῆ, ''gē'' ("earth") and -λoγία, ''-logia'', ("s ...
was primarily dry and warm, especially in arid areas. ''Proganochelys'' shared their environment with a variety of dinosaurs. ''Proganochelys'' lived in small water bodies such as ponds, but it was mainly earthbound.


References


External links


belated questions about beaks
{{Taxonbar, from=Q1193456 Monotypic prehistoric reptile genera Reptile taxonomy Norian first appearances Extant Late Triassic first appearances Late Triassic reptiles of Europe Late Triassic reptiles of Asia Prehistoric reptile genera Fossil taxa described in 1887 Taxa named by Georg Baur Testudinata