Low effective mechanical advantage of giraffes' limbs during walking reveals trade-off between limb length and locomotor performance.

Giraffes (Giraffa camelopardalis) possess specialized locomotor morphology, namely elongate and gracile distal limbs. While this contributes to their overall height and enhances feeding behavior, we propose that the combination of long limb segments and modest muscle lever arms results in low effective mechanical advantage (EMA, the ratio of in-lever to out-lever moment arms), when compared with other cursorial mammals. To test this, we used a combination of experimentally measured kinematics and ground reaction forces (GRFs), musculoskeletal modeling, and inverse dynamics to calculate giraffe forelimb EMA during walking. Giraffes walk with an EMA of 0.34 (±0.05 SD), with no evident association with speed within their walking gait. Giraffe EMA was about four times lower than expectations extrapolated from other mammals, ranging from 0.03 to 297 kg, and this provides further evidence that EMA plateaus or even diminishes in mammals exceeding horse size. We further tested the idea that limb segment length is a factor which determines EMA, by modeling the GRF and muscle moment arms in the extinct giraffid Sivatherium giganteum and the other extant giraffid, Okapia johnstoni. Giraffa and Okapia shared similar EMA, despite a four to sixfold difference in body mass (Okapia EMA = 0.38). In contrast, Sivatherium, sharing a similar body mass with Giraffa, had greater EMA (0.59), which we propose reflects behavioral differences, such as a somewhat increased capability for athletic performance. Our modeling approach suggests that limb length is a determinant of GRF moment arm magnitude and that unless muscle moment arms scale isometrically with limb length, tall mammals are prone to low EMA.

The locomotor kinematics and ground reaction forces of walking giraffes.

Giraffes (Giraffa camelopardalis) possess specialised anatomy. Their disproportionately elongate limbs and neck confer recognised feeding advantages, but little is known about how their morphology affects locomotor function. In this study, we examined the stride parameters and ground reaction forces from three adult giraffes in a zoological park, across a range of walking speeds. The patterns of GRFs during walking indicate that giraffes, similar to other mammalian quadrupeds, maintain a forelimb-biased weight distribution. The angular excursion of the neck has functional links with locomotor dynamics in giraffes, and was exaggerated at faster speeds. The horizontal accelerations of the neck and trunk were out of phase compared with the vertical accelerations, which were intermediate between in and out of phase. Despite possessing specialised morphology, giraffes' stride parameters were broadly predicted from dynamic similarity, facilitating the use of other quadrupedal locomotion models to generate testable hypotheses in giraffes.

The extinct, giant giraffid Sivatherium giganteum: skeletal reconstruction and body mass estimation.

Sivatherium giganteum is an extinct giraffid from the Plio-Pleistocene boundary of the Himalayan foothills. To date, there has been no rigorous skeletal reconstruction of this unusual mammal. Historical and contemporary accounts anecdotally state that Sivatherium rivalled the African elephant in terms of its body mass, but this statement has never been tested. Here, we present a three-dimensional composite skeletal reconstruction and calculate a representative body mass estimate for this species using a volumetric method. We find that the estimated adult body mass of 1246 kg (857-1812 kg range) does not approach that of an African elephant, but confirms that Sivatherium was certainly a large giraffid, and may have been the largest ruminant mammal that has ever existed. We contrast this volumetric estimate with a bivariate scaling estimate derived from Sivatherium's humeral circumference and find that there is a discrepancy between the two. The difference implies that the humeral circumference of Sivatherium is greater than expected for an animal of this size, and we speculate this may be linked to a cranial shift in centre of mass.

OSTEOCHONDROSIS IN THE DISTAL FEMURS OF AN ADULT RETICULATED GIRAFFE (GIRAFFA CAMELOPARDALIS RETICULATA): MACROSCOPIC, RADIOLOGIC, AND HISTOLOGIC FINDINGS.

An adult male reticulated giraffe (Giraffa camelopardalis reticulata) was presented for postmortem examination. During radiologic examination of the hindlimbs, osseous cyst-like lesions were detected in both medial femoral condyles. These lesions were subsequently examined macroscopically and histologically. The gross appearance suggested a diagnosis of bilateral osteochondrosis that was confirmed with histopathologic examination. This finding has not previously been reported in giraffes. Macroscopic visualization of the major limb joints, including the femorotibial joints, is therefore encouraged in future postmortem examinations of giraffes (Giraffa camelopardalis), and further assessment of clinical significance is required.

The running kinematics of free-roaming giraffes, measured using a low cost unmanned aerial vehicle (UAV).

The study of animal locomotion can be logistically challenging, especially in the case of large or unhandleable animals in uncontrolled environments. Here we demonstrate the utility of a low cost unmanned aerial vehicle (UAV) in measuring two-dimensional running kinematics from free-roaming giraffes (Giraffa camelopardalis giraffa) in the Free State Province, South Africa. We collected 120 Hz video of running giraffes, and calibrated each video frame using metatarsal length as a constant object of scale. We tested a number of methods to measure metatarsal length. The method with the least variation used close range photography and a trigonometric equation to spatially calibrate the still image, and derive metatarsal length. In the absence of this option, a spatially calibrated surface model of the study terrain was used to estimate topographical dimensions in video footage of interest. Data for the terrain models were collected using the same equipment, during the same study period. We subsequently validated the accuracy of the UAV method by comparing similar speed measurements of a human subject running on a treadmill, with treadmill speed. At 8 m focal distance we observed an error of 8% between the two measures of speed. This error was greater at a shorter focal distance, and when the subject was not in the central field of view. We recommend that future users maximise the camera focal distance, and keep the subject in the central field of view. The studied giraffes used a grounded rotary gallop with a speed range of 3.4-6.9 ms-1 (never cantering, trotting or pacing), and lower duty factors when compared with other cursorial quadrupeds. As this pattern might result in adverse increases in peak vertical limb forces with speed, it was notable to find that contralateral limbs became more in-phase with speed. Considering the latter pattern and the modest maximal speed of giraffes, we speculate that tissue safety factors are maintained within tolerable bounds this way. Furthermore, the angular kinematics of the neck were frequently isolated from the pitching of the body during running; this may be a result of the large mass of the head and neck. Further field experiments and biomechanical models are needed to robustly test these speculations.