Data and code from: Dipteran flight diversity is shaped by aerodynamic constraints, scaling, and evolutionary trade-offs
Data files
Jun 15, 2026 version files 2.47 GB
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Diptera_Aerodynamics.zip
294.30 KB
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Diptera_BodyMass_fresh_vs_dry.csv
15.70 KB
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Diptera_CFD.zip
231.48 MB
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Diptera_Explorer.zip
692.98 MB
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Diptera_Figure_Scripts.zip
40.54 KB
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Diptera_Kinematics_Aerodynamics.csv
6.17 KB
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Diptera_Kinematics_Coefficients_of_Variation.csv
1.28 KB
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Diptera_Kinematics.zip
518.13 KB
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Diptera_Morphology.csv
16.46 KB
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Diptera_Morphology.zip
746.83 KB
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Diptera_Photos.zip
1.49 GB
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Diptera_ThoraxMass.csv
1.77 KB
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Diptera_tree_family_level.RData
840 B
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Diptera_tree_species_level.RData
3.26 KB
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Diptera_Videos.zip
46.86 MB
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README.md
11.07 KB
Abstract
Flight has been a key innovation in insect evolution, yet the selective and mechanistic pressures shaping their flight motor systems remain poorly understood. Here, we present a comprehensive comparative analysis of flight in Diptera (true flies), integrating morphology, wingbeat kinematics, and aerodynamics within a phylogenetic framework. We quantified morphology in 133 species spanning the Dipteran phylogenetic and size range, and for a subset of 46 species we combined high-speed stereoscopic videography with computational fluid dynamics (CFD) to characterize wingbeat kinematics and aerodynamic performance, respectively. Our results reveal that morphology is strongly structured by phylogeny, whereas wingbeat kinematics are broadly conserved across Diptera, reflecting dominant aerodynamic constraints. Two early-diverged lineages, Culicomorpha (mosquitoes and midges) and Tipulomorpha (crane flies), exhibit strikingly divergent kinematics and aerodynamics, suggesting lineage-specific selective pressures. Combining these data with scaling analyses shows that maintaining in-flight weight support across the dipteran size range requires systematic allometric adjustments in wing morphology, wingbeat kinematics, and flight musculature. Smaller dipterans achieve weight support through relatively larger wings and higher wingbeat frequencies, whereas larger dipterans achieve the same aerodynamic requirement through increased investment in flight musculature to sustain the necessary mechanical power output. These size-dependent trait combinations highlight how different morphological and kinematic adaptations evolved in response to the shared physical requirements of hovering flight across Diptera. Mosquitoes and midges represent an extreme case, exhibiting a pronounced aerodynamic–acoustic trade-off with disproportionately large flight musculature and increased aerodynamic and acoustic power, consistent with selection favoring acoustic signaling during in-swarm mating. By integrating comparative morphology, kinematics, and aerodynamics across a major insect radiation, our study uncovers the interplay between physical scaling laws, aerodynamic constraints, and ecological pressures in shaping the evolution of animal flight. These findings provide a mechanistic framework for understanding how complex locomotor systems diversify under multiple selection pressures.
Dataset DOI: 10.5061/dryad.gxd25480s
Description of the data and file structure
Morphology, wingbeat kinematics, aerodynamics and phylogeny of the studied Diptera species.
Files and variables
File: Diptera_tree_species_level.RData
Description: Time-calibrated phylogenetic tree for the 133 Diptera species used in this study. The tree is pruned from Wiegmann & Yeates (2017) and formatted for use in comparative and phylogenetic analyses in R.
File: Diptera_tree_family_level.RData
Description: Time-calibrated phylogenetic tree for the 43 Diptera families studied. The tree is pruned from Wiegmann & Yeates (2017) and formatted for comparative and phylogenetic analyses in R.
File: Diptera_Morphology.csv
Description: Species-level mean morphological measurements for 133 Diptera species. Each row contains body and wing size, shape, and derived non-dimensional parameters.
Variables
- genus_species: species identifier (Genus_species format).
- suborder: suborder classification following Wiegmann & Yeates (2017).
- infraorder: infraorder classification following Wiegmann & Yeates (2017).
- family: family classification following Wiegmann & Yeates (2017).
- location: Sampling location (text description).
- flight kinematics: kinematics recorded (yes/no).
- body mass: Mean body mass. Units: mg.
- body length: Body length head-to-abdomen. Units: mm.
- body width: Maximum body width. Units: mm.
- body aspect-ratio: Ratio of body length to body width (non-dimensional).
- wing area: Total wing area. Units: mm².
- wing span: Distance between wing root and tip. Units: mm.
- wing mean chord: Mean chord length. Units: mm.
- wing aspect-ratio: Wing span² / wing area (non-dimensional).
- wing second-radius-of-gyration: Second moment-based radius of gyration. Units: mm.
- wing normalized second-radius-of-gyration: Second radius of gyration divided by wing span (non-dimensional).
File: Diptera_Kinematics_Aerodynamics.csv
Description: Species-level mean flight kinematics and aerodynamic parameters for 46 Diptera species. Each row contains wingbeat kinematics, derived aerodynamic forces, and non-dimensional metrics obtained from experimental measurements and aerodynamic modelling.
Variables
- genus_species: species identifier (Genus_species format).
- Body mass: Mean body mass. Units: mg.
- Wingspan: Distance between wing root and tip. Units: mm.
- Wing aspect-ratio: Ratio of body length to body width (non-dimensional).
- Wingspan-normalized second-radius-of-gyration: Second radius of gyration divided by wing span (non-dimensional).
- Wingstroke amplitude: Peak-to-peak stroke angle. Units: degrees.
- Wingbeat frequency: number of wingbeats per second. Units: Hz.
- Angle-of-attack: Mean aerodynamic angle of attack. Units: degrees.
- Reynolds number: Based on chord length and wing speed (non-dimensional).
- Strokeplane pitch angle: Angle of stroke plane relative to horizontal. Units: degrees.
- Lift coefficient: Force coefficient normal to the velocity vector (non-dimensional).
- Drag coefficient: Force coefficient tangential to the velocity vector (non-dimensional).
- Relative drag: Drag normalized by body weight (non-dimensional).
- Wing-tangential force coefficient: Force coefficient tangential to the wing surface (non-dimensional).
- Wing-normal force coefficient: Force coefficient normal to the wing surface (non-dimensional).
- relative tangential force : Tangential force normalized by body weight (non-dimensional).
- Weight-normalized aerodynamic power: Aerodynamic power divided by body weight. Units: W/N.
- Wingspan-normalized drag moment arm: Span-normalized moment arm of the drag force (non-dimensional).
- Wing speed: Wing speed at the drag moment arm. Units: m/s.
- Aeroacoustic power: Estimated wingbeat-induced sound power. Units: W.
- Normalized aeroacoustic power: Non-dimensional sound power.
File: Diptera_ThoraxMass.csv
Description: Species-level mean body mass and thorax mass measurements for 24 Diptera species. Each row includes total body mass, relative thorax mass, and the phylogenetically closest species in the flight dataset.
Variables
- genus_species: species identifier (Genus_species format).
- family: family classification following Wiegmann & Yeates (2017).
- infraorder: infraorder classification following Wiegmann & Yeates (2017).
- Body mass: Mean body mass. Units: mg.
- Relative thorax mass: fraction of total body mass (non-dimensional).
- closest species in flight dataset: phylogenetically closest species in the flight dataset.
File: Diptera_BodyMass_fresh_vs_dry.csv
Description: Paired fresh and dry body mass measurements for a subset of specimens. These data were used to estimate fresh body mass for specimens collected in French Guiana where only dry mass was available.
Variables
- specimen_ID: Unique identifier for each specimen.
- individual_ID: Identifier linking measurements from the same individual.
- capture_date: Date of specimen collection (DD.MM.YYYY).
- capture_place: Collection location (text description).
- GPS_coordinates (DDformat): Latitude and longitude in decimal degrees.
- fresh_mass_mg: Fresh body mass (mg).
- dry_mass_mg: Dried body mass (mg).
File: Diptera_Kinematics_Coefficients_of_Variation.csv
Description: Coefficients of variation (CV) for flight kinematics parameters, quantified within species (individual variation) and between species (interspecific variation).
Variables
- parameter: Name of the kinematic variable.
- variation_type: Type of variation: within-species or between-species.
- CV_value: Coefficient of variation (standard deviation divided by mean; non-dimensional).
File: Diptera_Morphology.zip
Description: Average wing outlines for 133 Diptera species used in the morphometric analysis. Each .csv file contains the mean wing contour of a single species, represented as a sequence of planar coordinates describing the wing boundary. Coordinates are ordered along the wing outline in the morphometrics coordinate system.
Variables
- X: spanwise coordinates of the wing outline. Units: normalized length (non-dimensional).
- Y: chordwise coordinates of the wing outline. Units: normalized length (non-dimensional).
File: Diptera_Kinematics.zip
Description: Time-resolved wingbeat kinematics for 46 Diptera species. Each .csv file contains the average wingbeat cycle of a single species, expressed as a time-normalized sequence of kinematic variables derived from flight measurements. The wingbeat is represented over a normalized time variable from 0 to 1, corresponding to one full stroke cycle.
Variables
- ft: Normalized time within the wingbeat cycle (0–1; non-dimensional).
- Stroke: Wing stroke angle within the stroke plane (degrees).
- Rotation: Wing rotation angle along the spanwise axis (degrees).
- Deviation: Wing deviation angle out of the stroke plane (degrees).
- Wingbeat frequency: Number of wingbeats per second (Hz).
- Stroke plane angle: Pitch angle of the stroke plane relative to horizontal (degrees).
File: Diptera_Aerodynamics.zip
Description: Time-resolved aerodynamic force coefficients for 46 Diptera species. The archive contains two folders:
- Force_Coeff_Lift_Drag: lift and drag force coefficients
- Force_Coeff_Normal_Tangential: force coefficients resolved normal and tangential to the wing stroke plane
The .csv files inside each folder contain the species-specific time-normalized sequence of aerodynamic force coefficients throughout the average wingbeat. All force coefficients are non-dimensional and derived from CFD simulations. The wingbeat is represented over a normalized time variable from 0 to 1, corresponding to one full stroke cycle.
Variables
- ft: Normalized time within the wingbeat cycle (0–1; non-dimensional).
- CL: Lift force coefficients normal to the wingbeat-induced velocity vector (non-dimensional).
- CD: Drag force coefficients parallel to the wingbeat-induced velocity vector (non-dimensional).
- CN: force coefficients normal to the wing surface (non-dimensional).
- CT: force coefficients tangential to the wing surface (non-dimensional).
File: Diptera_Photos.zip
Description: Microscope photographs of representative specimens for each of the 133 Diptera species. For each species, the dataset includes images of the full insect and an isolated wing, used for morphological characterization of wing shape data.
File: Diptera_Videos.zip
Description: High-speed videos of representative flying specimens for all 46 Diptera species studied in flight. For each species, the dataset includes a full-frame recording and a cropped, panning video focused on the flying insect.
File: Diptera_CFD.zip
Description: Parameter files for the CFD simulation of all 46 Diptera species studied in flight. Each folder contains:
- CFD input files (*.ini), required to re-run the simulation with our open-source code WABBIT (https://github.com/adaptive-cfd/WABBIT).
- CFD output files (*.t ), containing aerodynamic forces, moments, power, and related quantities used in the analysis.
All CFD data are dimensionless, as normalized with wingspan, wingbeat frequency, and density of air.
File: Diptera_Explorer.zip
Description: Interactive tool that enables phylogeny‑aware filtering, subset visualization, and direct data export. Users can select any family or species, which will highlight the corresponding wingbeat kinematics, aerodynamics, and underlying media (representative image and high-speed video). The export function allows side‑by‑side comparison of any insects in the dataset and complete reproduction of all figure‑level analyses.
Execution and requirements:
- Windows: run Diptera_Explorer.exe, which opens the application in the default web browser (tested in Chrome).
- Linux: run Diptera_Explorer.linux (may require execution permission via chmod +x).
- The application runs locally and requires a modern web browser.
- No installation or internet connection is required.
File: Diptera_Figure_Scripts.zip
Description: Custom R code for reproducing the manuscript figures from the provided data.
Code/software
- Data provided in CSV format can be loaded in most programming environments.
- The phylogenetic trees (.RData) can be loaded into R using: load(".../.RData").
- The R scripts can be executed in R using: source(".../*.R").
- CFD simulations can be reproduced using the open-source CFD code WABBIT (https://github.com/adaptive-cfd/WABBIT).
- The interactive tool runs locally via Diptera_Explorer.exe (Windows) or Diptera_Explorer.linux (Linux) and opens in a web browser; no installation or internet connection is required.
