26Plants producing toxic plant secondary metabolites (PSM) deter feeding of folivores. Animals 27 that are able to cope with noxious PSMs have a niche with a competitive advantage over other 28 species. However, the ability to cope with toxic PSMs incurs costs for detoxification. In order 29 to assess possible compensations for the ingestion of toxic PSMs, we compare the chemical 30 quality of plants consumed by bamboo lemurs (genera Hapalemur and Prolemur; 31 strepsirrhine primates of Madagascar) in areas with and without bamboo. Some bamboo 32 lemurs consume bamboo containing concentrations of cyanogenic substances 10 -50 times 33 above the average lethal dosage for mammals, and we postulate that animals consuming 34 Variation in food composition of bamboo lemurs 2 cyanogenic substances need supplementary protein or readily available energy for 35 detoxification. We compared the chemical composition of food consumed by three species of 36 bamboo lemurs that feed mainly (>80% of their time) on bamboo in the evergreen rainforest 37 of Ranomafana (Madagascar) with published data of the diets of bamboo lemurs at two sites 38 without highly cyanogenic plants (reed beds of Lac Alaotra and the evergreen littoral forest of 39 Mandena) and with food of sympatric folivorous lemur species that do not feed on bamboo. 40Lemurs feeding on bamboo consumed up to twice as much protein than bamboo lemurs in 41 areas without bamboo and sympatric lemur species that feed on leaves of trees. 42Concentrations of non-structural carbohydrates (a source of energy) showed the opposite 43 trend. This result supports the hypothesis that feeding on cyanogenic plants is linked to high 44 protein intake, either as a source of protein or for sulphur containing amino acids that can be 45 used for detoxification. Due to the high protein concentrations in bamboo, however, we 46 cannot distinguish between the hypothesis that lemurs that eat bamboo target additional food 47 items with higher protein, from the hypothesis that lemurs feeding on bamboo unavoidably 48 obtain higher concentrations of protein than animals feeding on leaves of trees, without an 49 added nutritional benefit. 50 51 54 1985; Nie et al. 2015), red panda (Ailurus fulgens; Johnson et al. 1988), pygmy rabbit 61 (Brachylagus idahoensis; Shipley et al. 2006), some marsupials (Dearing et al. 2000; Marsh et 62 al. 2003), and the bamboo lemurs of Madagascar (gentle and golden bamboo lemurs: 63 Hapalemur spp.; greater bamboo lemur: Prolemur simus; Mutschler Tan 1999Tan , 2006. 64Here, plants and folivorous animals are linked in an evolutionary arms race in which plants 65 produce PSM-to deter feeding, and folivores evolve morphological and/or physiological 66 mechanisms for detoxification of these components, or exhibit behavioural selection of higher 67 quality food or reduced energy expenditure to cope with PSMs (Freeland and Janzen 1974;