The origin of the turtle body plan: evidence from fossils and embryos

The origin of the unique body plan of turtles has long been one of the most intriguing mysteries in evolutionary morphology. Discoveries of several new stem‐turtles, together with insights from recent studies on the development of the shell in extant turtles, have provided crucial new information concerning this subject. It is now possible to develop a comprehensive scenario for the sequence of evolutionary changes leading to the formation of the turtle body plan within a phylogenetic framework and evaluate it in light of the ontogenetic development of the shell in extant turtles. The fossil record demonstrates that the evolution of the turtle shell took place over millions of years and involved a number of steps.

• Publication year

  2019

• Venue

  Palaeontology

• Publication date

  2019-12-02

• Fields of study

  Biology

• Identifiers
  DOI 10.1111/pala.12460
• External record

  Open on Semantic Scholar

• Source metadata

  Semantic Scholar

• No claims are published for this paper.

• No concepts are published for this paper.

• LSU Digital Commons LSU Digital Commons More than 1000 ultraconserved elements provide evidence that More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs turtles are the sister group of archosaurs

  2021cited by this paper
• Microanatomy of the stem-turtle Pappochelys rosinae indicates a predominantly fossorial mode of life and clarifies early steps in the evolution of the shell

  2019cited by this paper
• Postcranial remains of Teraterpeton hrynewichorum (Reptilia: Archosauromorpha) and the mosaic evolution of the saurian postcranial skeleton

  2019cited by this paper
• A Triassic stem turtle with an edentulous beak

  2018influential reference
• The early composition and evolution of the turtle shell (Reptilia, Testudinata)

  2018cited by this paper
• Osteology of the Middle Triassic stem-turtle Pappochelys rosinae and the early evolution of the turtle skeleton

  2018cited by this paper
• A Review of the Fossil Record of Basal Mesozoic Turtles

  2017cited by this paper
• Turtles as Hopeful Monsters: Origins and Evolution

  2017cited by this paper
• The amniote temporal roof and the diapsid origin of the turtle skull.

  2016influential reference
• Development of the turtle plastron, the order-defining skeletal structure

  2016cited by this paper
• Fossorial Origin of the Turtle Shell.

  2016cited by this paper
• Revision of the Triassic European turtles Proterochersis and Murrhardtia (Reptilia, Testudinata, Proterochersidae), with the description of new taxa from Poland and Germany

  2016influential reference
• A Middle Triassic palaeontological gold mine: The vertebrate deposits of Vellberg (Germany)

  2016cited by this paper
• Evolutionary origin of the turtle skull

  2015cited by this paper
• Neck motion in turtles and its relation to the shape of the temporal skull region

  2015cited by this paper
• A Middle Triassic stem-turtle and the evolution of the turtle body plan

  2015influential reference
• Emerging from the rib: resolving the turtle controversies.

  2015cited by this paper
• The origin of turtles: a paleontological perspective.

  2015influential reference
• Origin of the unique ventilatory apparatus of turtles

  2014cited by this paper
• Toward consilience in reptile phylogeny: miRNAs support an archosaur, not lepidosaur, affinity for turtles

  2014cited by this paper
• The first record of a nyctiphruretid parareptile from the Early Permian of North America, with a discussion of parareptilian temporal fenestration

  2014cited by this paper
• The hooked element in the pes of turtles (Testudines): a global approach to exploring primary and secondary homology

  2013cited by this paper
• Problems of the Ancestry of Turtles

  2013cited by this paper
• Anatomy, cranial ontogeny and phylogenetic relationships of the pareiasaur Deltavjatia rossicus from the Late Permian of central Russia

  2013cited by this paper
• Using Genes as Characters and a Parsimony Analysis to Explore the Phylogenetic Position of Turtles

  2013cited by this paper
• Biomechanics of turtle shells: how whole shells fail in compression.

  2013cited by this paper
• The Evolution of the Turtle Shell

  2013cited by this paper
• The ICS International Chronostratigraphic Chart

  2013cited by this paper
• The endoskeletal origin of the turtle carapace

  2013influential reference
• The draft genomes of soft–shell turtle and green sea turtle yield insights into the development and evolution of the turtle–specific body plan

  2013cited by this paper
• The girdles of the oldest fossil turtle, Proterochersis robusta, and the age of the turtle crown

  2013cited by this paper
• From Clone to Bone: Rocking clocks and clocking rocks: a critical look at divergence time estimation in mammals

  2012cited by this paper
• Phylogenomic analyses support the position of turtles as the sister group of birds and crocodiles (Archosauria)

  2012influential reference
• A developmental basis for innovative evolution of the turtle shell

  2012cited by this paper
• Reptilian-transcriptome v1.0, a glimpse in the brain transcriptome of five divergent Sauropsida lineages and the phylogenetic position of turtles

  2011cited by this paper
• Evolutionary developmental perspective for the origin of turtles: the folding theory for the shell based on the developmental nature of the carapacial ridge

  2011cited by this paper
• Homeotic effects, somitogenesis and the evolution of vertebral numbers in recent and fossil amniotes

  2010cited by this paper
• Transitional fossils and the origin of turtles

  2010influential reference
• The osteology and relationships of the Millerettidae (Reptilia: Cotylosauria)

  2009cited by this paper
• Proterochersis, eine pleurodire Schildkröte aus dem Keuper

  2009cited by this paper
• Timing of organogenesis support basal position of turtles in the amniote tree of life

  2009cited by this paper
• Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg

  2009cited by this paper
• Assembling the history of the Parareptilia: phylogeny, diversification, and a new definition of the clade

  2009cited by this paper
• A thin-shelled reptile from the Late Triassic of North America and the origin of the turtle shell

  2009cited by this paper
• An Archosaur-Like Laterosphenoid in Early Turtles (Reptilia: Pantestudines)

  2009cited by this paper
• Evolution of the Turtle Body Plan by the Folding and Creation of New Muscle Connections

  2009cited by this paper
• An ancestral turtle from the Late Triassic of southwestern China

  2008influential reference
• Development of the dermal skeleton in Alligator mississippiensis (Archosauria, Crocodylia) with comments on the homology of osteoderms

  2008cited by this paper
• Shell bone histology indicates terrestrial palaeoecology of basal turtles

  2007cited by this paper
• On the carapacial ridge in turtle embryos: its developmental origin, function and the chelonian body plan

  2007cited by this paper
• The Poverty of Taxonomic Characters

  2007cited by this paper
• The contribution of neural crest cells to the nuchal bone and plastron of the turtle shell.

  2007cited by this paper
• Calibration choice, rate smoothing, and the pattern of tetrapod diversification according to the long nuclear gene RAG-1.

  2007cited by this paper
• Anatomy and Relationships of Palaeochersis talampayensis, a Late Triassic Turtle from Argentina

  2007influential reference
• Functional morphology and evolution of aspiration breathing in tetrapods.

  2006cited by this paper
• Comprehensive survey of carapacial ridge‐specific genes in turtle implies co‐option of some regulatory genes in carapace evolution

  2005cited by this paper
• Integration of morphological data sets for phylogenetic analysis of Amniota: the importance of integumentary characters and increased taxonomic sampling.

  2005cited by this paper
• Sister group relationship of turtles to the bird-crocodilian clade revealed by nuclear DNA-coded proteins.

  2005cited by this paper
• DEVELOPING A PROTOCOL FOR THE CONVERSION OF RANK-BASED TAXON NAMES TO PHYLOGENETICALLY DEFINED CLADE NAMES, AS EXEMPLIFIED BY TURTLES

  2004cited by this paper
• A procolophonoid reptile with temporal fenestration from the Middle Triassic of Brazil

  2004influential reference
• Palaeoecology of Triassic stem turtles sheds new light on turtle origins

  2004cited by this paper
• Lung ventilation during treadmill locomotion in a terrestrial turtle, Terrapene carolina

  2003cited by this paper
• Owenetta kitchingorum, sp. nov., a small parareptile (Procolophonia: Owenettidae) from the Lower Triassic of South Africa

  2002cited by this paper
• Evidence for the neural crest origin of turtle plastron bones

  2001influential reference
• The evolutionary position of turtles revised

  2001cited by this paper
• Turtles as hopeful monsters.

  2001influential reference
• Morphogenesis of the turtle shell: the development of a novel structure in tetrapod evolution

  2001cited by this paper
• A molecular phylogeny of reptiles.

  1999cited by this paper
• THE ORIGIN AND EARLY EVOLUTION OF TURTLES

  1999cited by this paper
• Complete mitochondrial genome suggests diapsid affinities of turtles.

  1998cited by this paper
• Reptile phylogeny and the interrelationships of turtles

  1997cited by this paper
• Pareiasaur phylogeny and the origin of turtles

  1997cited by this paper
• A reassessment of Eunotosaurus africanus Seeley (Amniota: Parareptilia).

  1997cited by this paper
• Turtles as diapsid reptiles

  1996cited by this paper
• Correlated progression and the origin of turtles

  1996cited by this paper
• Late Triassic Turtles from South America

  1995cited by this paper
• A reevaluation of early amniote phylogeny

  1995cited by this paper
• The Origin of the Turtle Body Plan: Bridging a Famous Morphological Gap

  1993cited by this paper
• Studies on skeleton formation in reptiles: Patterns of ossification in the skeleton of Chelydra serpentina (Reptilia, Testudines)

  1993influential reference
• The Development and Evolution of the Turtle Body Plan: Inferring Intrinsic Aspects of the Evolutionary Process from Experimental Embryology

  1991cited by this paper
• The comparative osteology of the Triassic turtle Proganochelys. Bulletin of the AMNH ; no. 194

  1990influential reference
• Development of the turtle carapace: Implications for the evolution of a novel bauplan

  1989influential reference
• STAGES IN THE ORIGIN OF VERTEBRATES: ANALYSIS BY MEANS OF SCENARIOS

  1989cited by this paper
• AMNIOTE PHYLOGENY AND THE IMPORTANCE OF FOSSILS

  1988cited by this paper
• Vertebrate Paleontology and Evolution

  1988cited by this paper
• The Shell Morphology of the Triassic Turtle Proganochelys

  1985cited by this paper
• The Locomotion of Plesiosaurs

  1975cited by this paper
• Bauprinzipien im panzer der placodonten (reptilia triadica)

  1975cited by this paper
• Anatomy and function of expanded ribs in certain edentates and primates.

  1970cited by this paper
• THE PROBLEMATIC PERMIAN REPTILE EUNOTOSAURUS

  1969influential reference
• The mechanism of lung ventilation in the tortoise Testudo graeca Linné.

  1967cited by this paper
• Studies on the Structure & Development of Vertebrates

  1958cited by this paper
• BEITÄGE ZUR KENNTNIS DER ONTO‐GENIE UND DER VERGLEICHENDEN ANATOMIE DES SCHILDKRÖTEN‐PANZERS

  1942cited by this paper
• The homology of the shell elements in turtles

  1939cited by this paper
• STUDIES IN CHELONIAN OSTEOLOGY, Part II: THE MORPHOLOGICAL RELATIONSHIPS BETWEEN THE GIRDLES, RIBS AND CARAPACE

  1929cited by this paper
• The Phylogeny and Classification of Reptiles

  1917cited by this paper
• On the Classification of the Reptilia

  1916cited by this paper
• Die Wirbeltierfunde aus dem Keuper von Halberstadt

  1915cited by this paper
• Über die Phylogenie des Panzers der Schildkröten und über die Verwandtschaft der Lederschildkröte (Dermochelys coriacea)

  1914cited by this paper
• Über die Wirbeltierfunde in der oberen Trias von Halberstadt

  1914cited by this paper

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  2026cites this paper
• 3D shell asymmetry of Testudines as a potential biomarker for environmental stress

  2026cites this paper
• Investigation of Plasticity in Morphology, Organ Traits and Nutritional Composition in Chinese Soft-Shelled Turtle (Pelodiscus sinensis) Under Different Culturing Modes

  2025cites this paper
• Morphology and osteo-histology of the weigeltisaurid wing: Implications for aerial locomotion in the world's first gliding reptiles.

  2025cites this paper
• Evolution of the rostral vasculature in turtles

  2025cites this paper
• Insulin‐Like Growth Factor 1 ( IGF1 ) in the Chinese Soft‐Shelled Turtle, Pelodiscus sinensis : Molecular Characterization, Expression Pattern, and Polymorphism Analysis

  2025cites this paper
• First report of turtle and sauropod track assemblages from the Early Jurassic of Guizhou, China

  2025influential citation
• Stem Life: A Framework for Understanding the Prebiotic-Biotic Transition

  2024cites this paper
• Novel record of placodont remains including a Henodus cranium from the Upper Triassic Silves Group of the Algarve, southern Portugal

  2024cites this paper
• Functional and Character Disparity Are Decoupled in Turtle Mandibles

  2024cites this paper
• Thecal and Epithecal Ossifications of the Turtle Shell: Ontogenetic And Phylogenetic Aspects

  2024influential citation
• Growth and life history of freshwater chelydrid turtles (Testudines: Cryptodira): A bone histological approach

  2024cites this paper
• Multi-isotopic analysis reveals the early stem turtle Odontochelys as a nearshore herbivorous forager

  2023cites this paper
• Turtle skull development unveils a molecular basis for amniote cranial diversity

  2023cites this paper
• The topography of diet: Orientation patch count predicts diet in turtles

  2022cites this paper
• Constant 12 and Hahslm 472319 as Universe Creation Code on Tortoise Shell in Covid Economic Era

  2022cites this paper
• Gene Regulation during Carapacial Ridge Development of Mauremys reevesii: The Development of Carapacial Ridge, Ribs and Scutes

  2022cites this paper
• Correction to: Is the Shape of Turtle Claws Driven by Locomotor Modes?

  2022cites this paper
• Reptiles, Biodiversity of

  2022cites this paper
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  2022cites this paper
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  2021cites this paper
• Ontogenetic development of the European basal aquatic turtle Pleurosternon bullockii (Paracryptodira, Pleurosternidae)

  2021cites this paper
• Evolvability in the fossil record

  2021cites this paper
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  2021cites this paper
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  2021cites this paper
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  2021cites this paper
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  2021cites this paper
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  2020cites this paper
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  2020cites this paper
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  2020cites this paper
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  2020cites this paper