<i>Tetrapodophis amplectus</i>is not a snake: re-assessment of the osteology, phylogeny and functional morphology of an Early Cretaceous dolichosaurid lizard
Bibliographic record
Abstract
The origin of snakes remains one of the most contentious evolutionary transitions in vertebrate evolution. The discovery of snake fossils with well-formed hind limbs provided new insights into the phylogenetic and ecological origin of snakes. In 2015, a fossil from the Early Cretaceous Crato Formation of Brazil was described as the first known snake with fore- and hind limbs (Tetrapodophis amplectus), and was proposed to be fossorial, to exhibit large gape feeding adaptations (macrostomy) and to possess morphologies suggesting constriction behaviours. First-hand examination of T. amplectus, including its undescribed counterpart, provides new evidence refuting it as a snake. We find: a long rostrum; straight mandible; teeth not hooked zygosphenes/zygantra absent; neural arch and spines present and tall with apical epiphyses; rib heads not tubercular; synapophyses simple; and lymphapophyses absent. Claimed traits not preserved include: braincase/descensus parietalis; ‘L’-shaped nasals; intramandibular joint; replacement tooth crowns; haemal keels; tracheal rings; and large ventral scales. New observations include: elongate retroarticular process; apex of splenial terminating below posterior extent of tooth row; >10 cervicals with hypapophyses and articulating intercentra; haemapophyses with articulating arches; reduced articular surfaces on appendicular elements; rows of small body scales; and reduced mesopodial ossification. The axial skeleton is uniquely elongate and the tail with >100 vertebrae is not short as previously claimed, although overall the animal is small (∼195 mm total length). We assessed the relationships of Tetrapodophis using a revised version of the original morphological dataset, an independent morphological dataset, and these two datasets combined with molecular data. All four were analysed under parsimony and Bayesian inference and unambiguously recover Tetrapodophis as a dolichosaur. We find that Tetrapodophis shows aquatic adaptations and there is no evidence to support constricting behaviour or macrostomy.
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How this classification was reachedexpand
Full frame machine prediction
Teacher imitationNot calibrated prevalence, not ground truth. Human validation pending. The Gemma side is a direct model label for every work in the frame, read from the title-only record. The Codex side is a classifier learned from the 10,348 direct Codex labels and calibrated to design-weighted sample rates; fields without enough sample support carry no Codex call. Candidate is the union of the two sides; consensus is their intersection. These outputs are machine_predicted_unvalidated and are not human labels.
Distilled classifier scores by category (both heads)
| Category | Codex | Gemma |
|---|---|---|
| Metaresearch | 0.000 | 0.000 |
| Meta-epidemiology (narrow) | 0.000 | 0.000 |
| Meta-epidemiology (broad) | 0.000 | 0.000 |
| Bibliometrics | 0.001 | 0.000 |
| Science and technology studies | 0.001 | 0.001 |
| Scholarly communication | 0.001 | 0.001 |
| Open science | 0.000 | 0.001 |
| Research integrity | 0.000 | 0.000 |
| Insufficient payload (model declined to judge) | 0.003 | 0.001 |
Machine scores (provisional)
The two teacher heads of the student model, read on this work. A score orders the frame for review; it never asserts a category, and the validation status ships verbatim with every row.
Baseline scores from an immature model (maturity gate not passed, 7 training rounds). Scores rank; they never assert a category.
score_only:v0-immature-baseline · verbatim from the scoring run: score_only means the number may rank works, and no category label ships from itClassification
machine, unvalidatedMachine predicted; a candidate call from one source (direct Gemma or distilled Codex), not a consensus.
How this classification was reached, model by model and score by score, is at the end of the page under "How this classification was reached".