Physical limits of speed performances impose strong selective pressures on animals, influencing important functions such as predator avoidance and foraging success. Armoured herbivorous species such as tortoises represent a peculiar case: features that optimise encounter rate during reproduction, the ability to reach favourable sites for thermoregulation and resting, foraging or nesting sites may be more important than running speed. To explore this issue, we measured three characteristics related to travelling ability but that are independent of running speed: (1) muscular strength, (2) time needed for overturning and (3) jumping from a high step as proxies of the ability to overcome various obstacles. Additionally, reaction times in tortoises placed in a normal or overturned position were measured as a proxy of antipredator response. More than 400 adult Hermann’s tortoises from six populations were tested in the field during two seasons. Measures of travelling ability and antipredator response varied markedly among populations, and thus with environmental characteristics such as habitat type, terrain ruggedness and presence of predators. Tortoises from rugged and hot habitats (e.g. Mediterranean macchia) were the most successful and the fastest to accomplish the tests. Overturned tortoises were more reluctant to start moving compared to those in normal position, but this effect was absent in the two localities free from predators. Sex and season had limited effect on the measured performances. Overall, locomotor features essentially varied with environmental constraints. Future environmental studies should explore whether the observed differences among populations are linked to genetic adaptation or phenotypic plasticity.
Locomotor performances crucially influence habitat use, foraging and reproductive success, thus directly affecting individual fitness. Although running speed is often considered as a main indicator of agility, in armoured terrestrial vertebrates some other agility components might be more important. Heavy, rigid armour imposes trade-off between protection of soft body parts and locomotor performances and consequently habitat use. This study compiles three tests of locomotor performance which might be important for overcoming various obstacles present in mosaic habitats of tortoises. Testing adult Hermann’s tortoises from six populations with various habitat characteristics, we found significant inter-population differences. The ability of tortoises to complete the tests positively correlates with climatic conditions and topography of their habitats. Additionally, presence of predators in the habitats dramatically affects tortoise’s antipredator behaviour. Measured locomotor performances and antipredator behaviour showed surprisingly low level of sexual dimorphism.
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We are thankful to our colleagues Vuk Iković, Ana Vujović, Sonja Nikolić, Goran Šukalo, Rastko Ajtić and Gordan Kurbalija and his family for help during fieldwork. Miloš Popović helped with extracting ruggedness and climatic data. Research was partially financed by the Ministry of Education, Sciences and Technology of Republic of Serbia (grant no. 173043). We are thankful to the authorities for issuing the following permits: National Park “Galičica” (no. 11-4093/5), Agency for Environment Protection of Montenegro (no. UPI 2342/6) and Ministry of Environment, Mining and Spatial Planning of Serbia (no. 353-01-46/2012-03). We are thankful to two anonymous reviewers for helpful comments and suggestions.
Data availability statement
The datasets generated and analysed during the current study are available from the corresponding author on reasonable request.
This study was funded by the Ministry of Education, Sciences and Technology of Republic of Serbia (grant no. 173043).
Conflict of interest
The authors declare that they have no conflict of interest.
Experiments comply with the current laws of the countries in which they were performed. With the permissions of National Park “Galičica” (no. 11-4093/5), Agency for Environment Protection of Montenegro (no. UPI 2342/6) and Ministry of Environment, Mining and Spatial Planning of Serbia (no. 353-01-46/2012-03), animals were processed in the field. After non-invasive tests and measuring, which lasted up to 1 h, examinees were immediately released at the place of capture.
Communicated by T. Madsen
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Golubović, A., Anđelković, M., Arsovski, D. et al. Locomotor performances reflect habitat constraints in an armoured species. Behav Ecol Sociobiol 71, 93 (2017). https://doi.org/10.1007/s00265-017-2318-0
- Muscular strength
- Antipredator behaviour
- Hermann’s tortoise
- Testudo hermanni