Resources
Everything we built is free to use, change and redistribute under CC BY 4.0. That means credit us and the original data. Units are micrometres. Axes are +x anterior (head), +y dorsal (back) and −z the animal's left.
Cite both this project and the data it's built on; see how to cite. The organ names and the placement of the muscles are our identifications; the pages on the X-ray and muscles say how each was made and what's uncertain.
3D models
| File | What it is | Use it in |
|---|---|---|
| tardigrade-organs.glb 11.5 MB | Body surface and 19 organs and structures, one named mesh each | Web (three.js, model-viewer), Blender, Unity, Unreal |
| tardigrade-organs-stl.zip 13.8 MB | The same, one STL per structure | 3D printing, CAD |
| tardigrade-organs-obj.zip 7.3 MB | The same, one OBJ per structure | Blender, MeshLab, most 3D tools |
| tardigrade-muscles.glb 7.2 MB | 100 leg muscles (M1–M14, legs I–IV, both sides), placed on the X-ray animal, named like iiM5-L | Same as the organs; fits them in the same frame |
| tardigrade-muscles-stl.zip 8.8 MB | The same, one STL per muscle | 3D printing |
How this is built
- Data
- The nano-CT scans and organ label fields of Gross et al. 2019 (CC BY 4.0), and the confocal leg-muscle label fields of Gross & Mayer 2019 (Dryad, CC0).
- Built with
build_meshes.pyturns each organ label into a surface (marching cubes, then simplified) in micrometres.build_muscles.pynames each muscle from the paper's colour code and fits each leg onto the CT animal by two landmarks, foot and dorsal attachment, then nudges strands out of the organs.export_resources.pywrites the GLB, STL and OBJ files.- Checked
- By hand: the label files carry no names, so we matched each organ's measured volume to the paper's Table 1 and checked its position and number of pieces; uncertain ones are marked "probable". The muscle colours in each leg match Table 1's counts exactly (14, 12, 11 and 10).
- Limits
- The muscles come from a different animal and are placed approximately. Their slice spacing is estimated from round muscle strands, an assumption, and the right side is a mirror of the left.
- More
- How the whole project is built
Volumes and tables
| File | What it is | Use it in |
|---|---|---|
| tardigrade-labels-270nm.nrrd 0.5 MB | One label volume for the whole animal at 270 nm. It combines the published whole-body labels with the brain, salivary glands and head structures from the head scan, registered in, plus the traced body. Not available anywhere else. | 3D Slicer, napari, ITK-SNAP, Python (pynrrd) |
| tardigrade-labels-270nm.tif 0.6 MB | The same as a multi-page TIFF | Fiji / ImageJ |
| tardigrade-structures.csv | Label ids, names, confidence, measured and published volumes, and how each was identified | Anything |
| tardigrade-ct-bodymasked.nrrd 5.9 MB | The whole-body CT (8-bit), everything outside the traced body set to zero | Volume rendering, segmentation |
| tardigrade-gait.json | Every step of all eight legs in one recorded walk, group gait means (adults, juveniles, Ramazzottius, disulfiram), and the leg geometry on the X-ray animal. From Anderson et al. 2024. | Animation, robotics, gait analysis |
How this is built
- Data
- Gross et al. 2019's whole-body scan and labels at 270 nm and their head scan's labels; walking data from Anderson et al. 2024's repository (no licence stated).
- Built with
export_resources.pymerges the whole-body labels, our traced body and the head-scan labels. We placed the head scan by cross-correlating the two scans over all 24 orientations, then refined the shift on the pharynx, buccal tube and ventral ganglion.export_gait.pyturns the walking workbooks into step events and group means.- Checked
- The head scan's stated voxel size did not fit its data. At the size we measured, 0.1065 µm, its brain and salivary glands come to 1,389 and 2,274 µm³, against the published 1,393 and 2,274.
- Limits
- The head-scan voxel size is our measurement, not the paper's figure. The walk is one recorded animal plus group means.
- More
- How the whole project is built
Predicted protein structures
Eleven co-folding predictions from the condensin II work: each tardigrade CAP-H2 kleisin region placed on its own species' SMC head, with the human pair as a positive control. Each file is the best-scoring sample of several.
These are computed models, not experimental structures. ipTM is the confidence in the interface between the two chains and pLDDT the per-residue confidence of the kleisin; scrambled-sequence controls scored 0.19–0.59, so treat anything below about 0.6 as unsupported. They cannot be deposited in the PDB, which takes experimental structures only — ModelArchive is where computed models belong and is where these are headed.
| File | What it is | Confidence |
|---|---|---|
| c_pme_a.pdb 435 kB | P. metropolitanus — Paramacrobiotus metropolitanus CAP-H2 copy a, C-terminal region, on its own SMC4 head | ipTM 0.808, pLDDT 62 |
| c_pme_b.pdb 426 kB | P. metropolitanus — Paramacrobiotus metropolitanus CAP-H2 copy b, C-terminal region, on its own SMC4 head | ipTM 0.805, pLDDT 55 |
| c_hex_b.pdb 408 kB | H. exemplaris — Hypsibius exemplaris CAP-H2 copy b, C-terminal region, on its own SMC4 head | ipTM 0.794, pLDDT 66 |
| c_hex_a.pdb 412 kB | H. exemplaris — Hypsibius exemplaris CAP-H2 copy a, C-terminal region, on its own SMC4 head | ipTM 0.777, pLDDT 50 |
| c_rva_b.pdb 413 kB | R. varieornatus — Ramazzottius varieornatus CAP-H2 copy b, C-terminal region, on its own SMC4 head | ipTM 0.763, pLDDT 63 |
| c_rva_a.pdb 430 kB | R. varieornatus — Ramazzottius varieornatus CAP-H2 copy a, C-terminal region, on its own SMC4 head | ipTM 0.677, pLDDT 38 |
| n2_hex_b.pdb 452 kB | H. exemplaris — Hypsibius exemplaris CAP-H2 copy b, N-terminal region, on its own SMC2 head and neck | ipTM 0.665, pLDDT 68 |
| n2_rva_b.pdb 451 kB | R. varieornatus — Ramazzottius varieornatus CAP-H2 copy b, N-terminal region, on its own SMC2 head and neck | ipTM 0.655, pLDDT 64 |
| n2_pme_b.pdb 443 kB | P. metropolitanus — Paramacrobiotus metropolitanus CAP-H2 copy b, N-terminal region, on its own SMC2 head and neck | ipTM 0.415, pLDDT 44 |
| c_hsa.pdb 400 kB control | Homo sapiens — Human CAP-H2 C-terminal region on human SMC4 head, as the positive control | ipTM 0.834, pLDDT 74 |
| n2_hsa.pdb 451 kB control | Homo sapiens — Human CAP-H2 N-terminal region on human SMC2 head and neck, as the positive control | ipTM 0.613, pLDDT 58 |
Method and the full sample set: the condensin II preprint, doi:10.5281/zenodo.22832510. Predicted with Boltz-2.
Minecraft
| File | What it is | Use it in |
|---|---|---|
| tardigrade_270nm.schem 0.3 MB | Glass skin and concrete organs, 608 × 188 × 213 blocks | Java 1.20+ with WorldEdit, FAWE or Litematica |
| tardigrade_ct_270nm.schem 2.1 MB | The X-ray itself in 11 grey stones, solid | Same |
| README.txt | How to load them, block legend, credits |
How this is built
- Data
- The same nano-CT and labels (Gross et al. 2019, CC BY 4.0), with the head structures placed by our registration.
- Built with
export_minecraft.pyturns one 270 nm voxel into one block and writes Sponge schematics: a one-block glass skin with concrete organs, and a second build where each voxel's CT brightness picks one of 11 grey blocks.- Checked
- Not tested against data; it is a direct conversion of the scan.
- Limits
- One block stands for 270 nm, so the animal is about 3.7 million times life size and carries the scan's resolution, no finer.
- More
- How the whole project is built
The knowledge graph
Everything the project knows, as 2,500 linked facts, sources, open questions, tests and assumptions, each citing the page or record it came from. Search it in the browser, or download nodes.jsonl.gz and edges.jsonl.gz. CC BY 4.0.
Videos
Walking, the block build and the stone build, each wide, square and tall: on the front page. Share freely with credit.
Tools
We wrote these along the way. Each works beyond tardigrades. The code is on GitHub: tardigraph-tools (MIT licence), and the condensin analysis has its own repository.
| Tool | What it does |
|---|---|
| Label fields to meshes | Turns Amira and ImageJ label volumes into named, simplified meshes (GLB, STL, OBJ). Traces a body surface from a CT cuticle, handling mounting film. |
| Scan-to-scan registration | Places a high-resolution partial scan inside a whole-body scan: FFT cross-correlation over all 24 orientations, then refinement on label overlap. |
| Muscle placement | Fits segmented structures from one specimen onto another by landmarks, then nudges them out of the target's organs. |
| Voxels to Minecraft | Writes Sponge schematics from any label or greyscale volume, with no dependencies beyond NumPy. |
| Browser viewer | A three.js viewer for organ meshes: glass body, CT on the cut face, grouped structure list, walking from step events, shareable view links. |
| Video recorder | Renders frame-exact videos of browser 3D scenes in wide, square and tall, on the GPU. |
| Reciprocal ortholog check | Tests whether "missing" genes are really missing: reciprocal best hits and PSI-BLAST across related proteomes. |
| Remote tar indexing | Finds files inside huge remote tar archives (such as OpenGenome2) with HTTP range requests, without downloading them. |
| Methylation from HiFi BAMs | Reads 5mC calls (MM/ML tags) from public PacBio files, with checks for amplified libraries. |
Preprints
Everything we have written up, with its data and the script that produced it. All CC BY 4.0, all open, all with their failures and corrections in the text rather than trimmed out of it.
- Tardigrades have a complete condensin II: two divergent CAP-H2 kleisins in three eutardigrade genomes. Version 4, with public errata. doi:10.5281/zenodo.22832510 (all versions) · code and data
- Condensin II never switches on in a tardigrade embryo: it is maternally supplied and shuts down with the division machinery. The functional half of the story above, from 62 single timestamped embryos — including the confound test we had to replace because the first one was circular, and two corrections to what we published the day before. doi:10.5281/zenodo.22879340
- Seed at cell centres, not nuclei: the cost of a standard shortcut in tessellation models of embryos. Where a nucleus sits inside its own cell (60,701 cell-frames), what seeding a model there costs, and why neither obvious correction recovers it. Version 2 (6 October 2026) corrects one sentence: Guignard et al. 2020 had already measured contact-area variability across ascidian embryos, and we said nobody had. doi:10.5281/zenodo.22879282 (all versions)
- A tardigrade builds its synapses last: two developmental programmes in Hypsibius exemplaris, separated by their timing and then by their chemistry When a tardigrade embryo transcribes its synapses, from two public transcriptomes nobody had asked: the synaptic genes rise last and fall after hatching, the patterning genes pulse early, and an unc-86 orthologue switches on inside the pulse. With the test that failed, the one that rescued it, and a time axis we had wrong by a factor of two. doi:10.5281/zenodo.23163786
- A conserved achaete-scute cluster in three tardigrade genomes, invisible to all three annotations Two achaete-scute genes side by side in all three annotated tardigrade genome, found by searching the sequence after the annotations said they were not there. doi:10.5281/zenodo.23163792
- The two camps on the tardigrade brain have been looking at different animals: a structured reading of the head-segment debate Every claim in the forty-year argument over the tardigrade brain, laid out by the species it was made on, and a widely cited table that has Kristensen's 1982 protocerebrum backwards. Version 2 (6 October 2026) adds Schulze et al. 2014, the one species both camps studied, with opposite conclusions. doi:10.5281/zenodo.23163804 (all versions)
- A molecular tree of 412 named tardigrade species from all public 18S, 28S and COI, with the records it cannot accept. A species-level tree of the phylum from every public sequence that matches a checklist name, three times the 128 named species in the last phylum-wide tree (Guil et al. 2019), and the screen that threw out 176 records with written evidence. Every conflict with the checklist checked against the literature: none is new. With the tree, alignments, screen and script. doi:10.5281/zenodo.23184587
- What is known about the tardigrade nervous system, and in which animal it was seen: a per-structure concordance, with the first measurements of the trunk ganglia. Every published statement about each part of the nervous system, with the species it was actually seen in: 42% are from the laboratory animal, and 38 of 104 structures have never been described in it. With the full concordance as a table and as JSON. doi:10.5281/zenodo.23164252
How to cite
Please cite this project, and the data you used.
Meow-Ludo (2026). Tardigraph: a water bear rebuilt from published data. https://tardigraph.meow-meow.io/ (accessed [date]). CC BY 4.0.
@misc{tardigraph2026,
author = {Meow-Ludo},
title = {Tardigraph: a water bear rebuilt from published data},
year = {2026},
url = {https://tardigraph.meow-meow.io/},
note = {CC BY 4.0}
}
And the original data:
- Organs, CT, blocks: Gross V, Müller M, Hehn L, Ferstl S, Allner S, Dierolf M, Achterhold K, Mayer G, Pfeiffer F (2019). X-ray imaging of a water bear offers a new look at tardigrade internal anatomy. Zoological Letters 5:14. doi:10.1186/s40851-019-0130-6
- Muscles: Gross V, Mayer G (2019). Cellular morphology of leg musculature in the water bear Hypsibius exemplaris (Tardigrada) unravels serial homologies. R Soc Open Sci 6:191159. doi:10.1098/rsos.191159
- Walking: Anderson EM et al. (2024). Comparative analysis of tardigrade locomotion across life stage, species, and disulfiram treatment. PLOS ONE. doi:10.1371/journal.pone.0310738
Full list: credits.