Fecal chlorophyll describes the link between primary production and consumption in a terrestrial herbivore

Spatiotemporal variation in primary productivity is known to have strong and far-reaching effects on herbivore ecology, but this relationship is often studied indirectly at broad scales, in part due to the difficulty in measuring selection for green biomass by individual animals. In aquatic systems, the concentration of chlorophyll in herbivore feces has been used as a direct measure of the consumption of photosynthetic primary production, but this method has not been applied to terrestrial systems. We measured chlorophyll concentration in feces from elk (Cervus elaphus) experiencing large fluctuations in primary production in the winter to spring transition over three years. We compared temporal trends in fecal chlorophyll with trends in fecal nitrogen, grass chlorophyll, grass digestible nitrogen, and landscape-level primary productivity (as described by the normalized difference vegetation index or NDVI). We also directly examined the relationship between fecal chlorophyll and NDVI. Temporal trends in fecal chlorophyll were strong and well described by piecewise regression (adjusted coefficient of determination, r(2)a = 0.881-0.888), showing uniformly low concentrations throughout winter followed by an abrupt, rapid increase beginning on different Julian days (88, 91, or 110) each year. Changes in fecal chlorophyll closely matched the temporal trend in the chlorophyll and digestible nitrogen concentration of forage grasses collected directly from elk feeding sites. Fecal chlorophyll also tracked broad temporal patterns in fecal nitrogen and NDVI, but discrepancies between the indexes may highlight preferences or constraints on selectivity for green biomass in elk. Spatially and temporally matched NDVI and fecal chlorophyll estimates were uncorrelated until NDVI reached approximately half its seasonal range. Combined, these data describe important patterns in selection for nutritious, green biomass in a temperate herbivore that would be difficult to study without data on fecal chlorophyll. Fecal chlorophyll produced novel and precise descriptions of (and detected large interannual differences in) winter length, severity, and the rate of spring green-up, as they were experienced by a large, grazing herbivore. Measuring fecal chlorophyll provides a noninvasive, inexpensive, and direct approach to describe an important aspect of foraging ecology in terrestrial herbivores and may be particularly powerful for studying climate effects in seasonal environments.