Article: steppe tortoise diet in the wild

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GeoTerraTestudo

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Diet of the steppe tortoise at the Djeiron Ecocenter of Bukhara, Republic of Uzbekistan, near the Kyzyl Kum Desert (Lagarde et al. 2003a).

Steppe tortoises were active for only 3 months out of the year, with males emerging from brumation in March and going back into aestivation in May, and females emerging in April and returning in June. There was some sporadic activity in the autumn (September-October), but mostly the tortoises aestivated from June-November, and then brumated from November-March. Mating season was March-April, and the post-mating season was April-May. Diet was determined both by plant availability and by preference.

Plants consumed during the mating season (in order of decreasing % fresh mass consumed):

1) Ceratocephalus (Ranunculaceae, or buttercup family) - 64.84%
2) Centaurea (Asteraceae, or daisy family) - 19.87%
3) Epilasia (Asteraceae, or daisy family) - 4.86%
4) Koelpinia (Asteraceae, or daisy family) - 4.75%
5) Brassicaceae sp. (cabbage family) - 4.56%
6) Asteraceae sp. (daisy family) - 0.43%
7) Veronica (Plantaginaceae, or plantain family) - 0.32%
8) Papaveraceae sp. (poppy family) - 0.23%
9) Hypecoum (Papaveraceae, or poppy family) - 0.07%
10) Alyssum (Brassicaceae, or cabbage family) - 0.03%

Plants consumed during the post-mating season (in order of decreasing % fresh mass consumed):

1) Papaver (Papaveraceae, or poppy family) - 36.51%
2) Koelpinia (Asteraceae, or daisy family) - 25.59%
3) Brassicaceae sp. (cabbage family) - 20.73%
4) Asteraceae sp. (daisy family) - 5.53%
5) Epilasia (Asteraceae, or daisy family) - 4.73%
6) Ceratocephalus (Ranunculaceae, or buttercup family) - 3.34%
7) Roemaria (Papaveraceae, or poppy family) - 1.72%
8) Centaurea (Asteraceae, or daisy family) - 1.17%
9) Hypecoum (Papaveraceae, or poppy family) - 0.36%
10) Veronica (Plantaginaceae, or plantain family) - 0.26%

Based on this study, it looks like the only plants favored in both periods were in the daisy family. The other plants dominating the diet were buttercups, poppies, and Brassicaceae (cabbage family).
 

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GeoTerraTestudo

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Also should mention that all of these plants were annuals. The article also said that in the wild, steppe tortoises do not compete with ungulates. This is because sheep, goats, cattle, and horses ate grass and other weeds that the tortoises did not prefer. Also, the tortoises were able to eat plants like the buttercup (Ranunculaceae) that would sicken livestock or even kill them if eaten in large enough quantities, while the tortoises were able to eat them with no ill effects.

There are some indications, though, that steppe tortoises eat more toxic weeds when they have a high gut parasite load, and that they eat milder weeds when they have a more normal gut biota. In other words, it appears that the tortoises are self-medicating by eating bitter- or acrid-tasting plants to deworm themselves, and then eating tastier plants when they are healthier.
 
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