Dg. Williams et Jr. Ehleringer, Intra- and interspecific variation for summer precipitation use in pinyon-juniper woodlands, ECOL MONOGR, 70(4), 2000, pp. 517-537
In the arid southwest of North America, winter precipitation penetrates to
deep soil layers, whereas summer "monsoon" precipitation generally wets onl
y surface layers. Use of these spatially separated water sources was determ
ined for three dominant tree species of the pinyon-juniper ecosystem at six
sites along a gradient of increasing summer precipitation in Utah and Ariz
ona. Mean summer precipitation ranged from 79 to 286 mm, or from 18% to 60%
of the annual total across the gradient. We predicted that, along this sum
mer rainfall gradient, populations of dominant tree species would exhibit a
clinal off-on response for use of water from upper soil layers, responding
at particular threshold levels of summer precipitation input. This predict
ion was largely supported by our observations of tree water source use over
a two-year period and from irrigation experiments.
Hydrogen and oxygen stable isotope ratios (deltaD and delta O-18) of tree x
ylem water were compared to that of precipitation, groundwater, and deep an
d shallow soil water to distinguish among possible tree water sources. delt
aD-delta O-18 relationships and seasonal xylem water potential changes reve
aled that trees of this ecosystem used a mixture of soil water and recent p
recipitation, but not groundwater. During the monsoon period, a large propo
rtion of xylem water in Pinus edulis and Juniperus osteosperma was from mon
soon precipitation, but use of this precipitation declined sharply with dec
reasing summer rain input at sites near the regional monsoon boundary in Ut
ah. Quercus gambelii at most sites along the gradient used only deep soil w
ater even following substantial inputs of summer rain. Populations of Querc
us at sites with the highest average summer precipitation input, however, p
redominantly used water in upper soil layers from recent summer rain events
. Soil temperature correlated with patterns of summer precipitation use acr
oss the gradient; high soil temperatures north of the monsoon boundary may
have inhibited surface root activity for some or all of the three tree spec
ies.
Irrigation experiments with deuterium-labeled water revealed that Quercus g
ambelii in northern Arizona and southern Utah did not use water from surfac
e layers. In contrast, Juniperus osteosperma at these sites responded signi
ficantly to the irrigations: between 37% and 41% of xylem water originated
from irrigations that wetted only the top 30 cm of soil. Responses by Pinus
edulis to these irrigations were variable; uptake of labeled water by this
species was greater in September at the end of the summer than during the
hot midsummer period. Inactivity of Pinus roots in midsummer supports the h
ypothesis that root activity in this species is sensitive to soil temperatu
re.
Seasonal patterns of leaf gas exchange and plant water potential correspond
ed to the seasonality of rainfall at different sites. However, no correlati
on between a species' ability to use summer rainfall and its tolerance to w
ater deficits at the leaf level was found. Midday stomatal conductance (g(s
)) for Pinus needles approached zero at predawn water potentials near -2 MP
a, whereas g(s) in Quercus and Juniperus declined to zero at -2.8 and -3.7
MPa, respectively. The relationship between photosynthesis (A) and g(s) was
similar among the three species, although Quercus maintained higher overal
l rates of gas exchange and tended to operate higher on the A/g(s) curve th
an the two conifers. At sites in eastern Arizona where Quercus fully used m
oisture from summer rains, leaf gas exchange characteristics were similar t
o those of Pinus and Juniperus.