Mat
muhly (Muhlenbergia richardsonis),
at almost 4000 meters in eastern California's White Mountains, holds the
highest elevation record for a C4 species in North America. Archival herbaria
and climate data indicate this species has been moving upslope in the last
10-20 years, apparently as a result of recent climate change. A comparative
study of foliar characteristics was done along a 3000-3800 meter elevational
gradient with M. richardsonis
and three co-occurring C3 graminoid species. In all species, we examined
elevation-dependent variation in stomatal density (SD), leaf nitrogen content
(%N), and leaf tissue d 13C. These foliar traits generally vary with
elevation in C3 species but have not been examined across elevation in C4
species. Leaf SD, %N, and d 13C all increased with elevation as expected in
the C3 plants. Leaf SD and d 13C were unchanging with elevation in M.
richardsonis but %N increased
with elevation as it did in the C3 species. We interpret the C3/C4 contrasts in
SD and d 13C data to mean that C4 photosynthesis is not
limited by the low partial pressures of atmospheric CO2 (pCO2) present at high
elevations. C3 plants are limited by low pCO2 at high elevation and shifts in
SD and d 13C reflect compensatory responses to cope with
the thinning atmosphere. The similar trend in %N for C3 and C4 tissues suggests
this is a response to altitudinal variables other than pCO2 (e.g. temperature
or precipitation regimes) that affects C3 and C4 plants similarly. C4 plants
are rare at high elevations for reasons that are not well understood. This
study indicates C4 plants are not excluded from high elevations by low pCO2.
Rather, in this respect, C4 plants appear to be pre-adapted to the thin
atmospheres of alpine habitats.