A giraffe’s hoof must do more than resist wear. It has to remain attached to living tissue, accommodate continued growth and transmit the forces of a heavy animal taking a step. A uniformly hard block would tell only part of that story. The arrangement of the softer structures inside it matters too.
Liza Dadone and colleagues examined that arrangement in feet collected from six free-ranging southern giraffes. Their 2025 anatomical study combined dissection with microscopic sections to connect the visible shape of the hoof with the tissues beneath it. They found clear regional differences: the centre of the sole, the toe and the heel were built differently, and the internal cushion itself contained two contrasting materials.
The work addresses a basic difficulty in interpreting giraffe feet. An unusual image is easier to recognise as abnormal when the normal structures have first been described. The study supplies that comparison at a tissue level, complementing research that uses radiographs or CT to examine bones and lesions.
A shell maintained by living tissue
The hoof capsule is the outer covering made of keratinised tissue, or horn. Beneath it lies the corium, a layer of living connective tissue with a blood supply. The distinction is important: the epidermal cells make the horn, while the corium supports those cells and provides the structures around which the horn is organised.
Some parts of this boundary form finger-like projections called papillae. Elsewhere, the tissues interlock through sheets called laminae. Both increase contact between the living foot and its covering, but they are arranged differently according to their location. The laminar region helps anchor the hoof wall; it is not simply another identical horn-producing surface.
The team could see these relationships by removing the hoof capsule from a front and a hind foot of one adult male. They also cut lengthwise through the inner digit of the right front foot from each of the six animals. These sections put the outer hoof, the last toe bone, the tendon and the cushion into the same anatomical view. Samples from selected regions were then examined under the microscope.
The animals—four males and two females—had already been killed in a reserve management cull before the research project was conceived. Their feet were collected opportunistically and frozen. They were adults of unknown exact ages, so the study describes a small adult sample rather than a developmental series or a population-wide reference range.
The sole has a centre and a rim
Viewed from below, the hoof had a raised peripheral wall around a concave central sole. In the lengthwise sections, the horn was thickest beneath the toe tip and heel and thinner through the centre. These were relative anatomical comparisons visible in the sections, not a standardized numerical survey of sole thickness.
The tissue beneath the horn also changed with location. Papillae near the toe and wall margins were longer and more robust than those in the central sole. Along the front-facing surface of the last toe bone, the upper region had papillae while the lower region contained long laminae. The microscopic sections showed the same regional organisation seen during dissection.
This pattern fits a hoof whose different regions perform different mechanical tasks. More substantial horn and supporting tissue occur around surfaces expected to meet the ground early and sustain loading. But the researchers did not measure pressure beneath a walking giraffe. The anatomy supports a functional interpretation; it does not provide a map of force through the entire step.
An earlier study of wild Nubian giraffes in Uganda offers a useful comparison. Dadone, Steve Foxworth, Matthew Johnston and their collaborators pressed a painted board against the undersides of feet while the giraffes were lying down during translocation procedures. Paint marked prominent peripheral surfaces, especially the hoof wall and heel, while the recessed centre remained unpainted.
That simple observation independently established the initial contact geometry. It did not mean the central sole never bears weight: a foot deforms when the animal loads it. The later tissue study helps explain what lies beneath that geometry, in a different giraffe population and with a different method.
A cushion made from two materials
The digital cushion sits inside the foot near the heel, behind the deep tendon that flexes the digit. Its name can suggest a single pad, but the dissections revealed two regions with distinct textures.
The upper portion was soft and rich in fat. The lower portion, extending toward the sole, was firmer and consisted of dense fibrous connective tissue. Microscopy confirmed that the difference was in the tissues themselves, rather than merely their appearance in a cut specimen.
A combination of deformable and stronger fibrous material offers a plausible way to spread forces while supporting nearby structures. The authors proposed roles in cushioning, protecting the tendon and maintaining circulation through the foot. Those functions remain hypotheses for direct mechanical and physiological investigation in giraffes; identifying the compartments does not measure how much shock either absorbs.
The distinction nevertheless matters when interpreting a scan or a tissue sample. Normal fat and dense connective tissue should look different. A future comparison of healthy and altered cushions will need to compare the corresponding regions, rather than treating any difference within the cushion as damage.
A reference with natural variation
Most sampled hooves followed a similar relationship between the sole and the lower surface of the last toe bone. One showed a localized bulge of excess sole toward the heel. Even in a free-ranging group, the external shape was not perfectly uniform.
The examined tissue sections did not reveal the significant bone, tendon or supporting-tissue lesions the researchers screened for. That finding supports using the material as an anatomical reference, but it is not proof that every part of each foot was lesion-free. Only selected regions were sectioned, and the study did not include advanced imaging of the entire foot. Freezing and thawing also complicated interpretation of colour and apparent bleeding.
These constraints define the next useful comparisons. Examining matching regions from altered hooves could show which tissues change as the capsule changes shape. Measuring pressure during movement could test whether the regional architecture corresponds to the proposed distribution of forces. Neither question can be answered by external shape alone.
The study makes the giraffe hoof more intelligible as a connected structure: growing horn follows an organised living boundary, the sole varies across its surface, and the internal cushion combines different materials. Understanding those relationships gives subsequent investigations something concrete to compare when the structure or function of a foot changes.
Sources
- Liza Dadone, Steve Foxworth, Jacqueline Goedhals, Sushan Han, Priya Bapodra-Villaverde, Seng Wai Yap, Thato Radile and Francois Deacon (2025). “Anatomy and baseline histology of the hoof capsule, corium, and digital cushion in free-ranging southern giraffe (Giraffa giraffa).” PLOS ONE 20, e0339972. Full paper.
- Liza Dadone, Steve Foxworth, Robert Aruho, Amy Schilz, Andrea Joyet, Myra Barrett, Peter Morkel, Garrett Crooks, Julian Fennessy and Matthew S. Johnston (2021). “Foot shape and radiographs of free-ranging Nubian giraffe in Uganda.” PLOS ONE 16, e0252929. Related study.
