Scaling and foraging behavior drive the evolution of humeral shape in hummingbirds
Bibliographic record
Abstract
Abstract Understanding how locomotion-related skeletal elements evolve under biomechanical and ecological constraints is central to animal evolutionary biology. In hummingbirds (Trochilidae), the humerus plays a key role in force transmission during hovering and flapping flight, yet the drivers of its shape evolution have not been examined. We combined geometric morphometrics with phylogenetic comparative analyses to examine humeral shape variation, evolutionary rates, and phenotypic integration in male hummingbirds from 78 species. Our analyses identified humerus allometry as the dominant predictor of shape, revealing a pattern in which larger humeri show broader proximal epiphyses, increased shaft robustness, and reduced curvature. In addition to this scaling pattern, we find that male humerus shape differs subtly among hummingbird species that differ in the use of aggression during nectar foraging. Evolutionary rates of humeral shape were heterogeneous and decoupled from ecological predictors. We also find phenotypic integration between the proximal and distal regions of the humerus, indicating coordinated evolution. Together, these results show that humeral evolution in hummingbirds is governed primarily by biomechanical scaling and internal integration, with foraging ecology introducing secondary, size-dependent modifications. This work highlights the importance of considering scaling and internal integration when interpreting morphological evolution in locomotor systems across vertebrates.
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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 | 0.001 |
| Meta-epidemiology (narrow) | 0.000 | 0.000 |
| Meta-epidemiology (broad) | 0.000 | 0.000 |
| Bibliometrics | 0.001 | 0.000 |
| Science and technology studies | 0.000 | 0.000 |
| Scholarly communication | 0.000 | 0.000 |
| Open science | 0.000 | 0.000 |
| Research integrity | 0.000 | 0.000 |
| Insufficient payload (model declined to judge) | 0.002 | 0.000 |
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".