Yellowstone's Northern Range - Greater Yellowstone Science ...
Yellowstone's Northern Range - Greater Yellowstone Science ...
Yellowstone's Northern Range - Greater Yellowstone Science ...
Create successful ePaper yourself
Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.
WOODY VEGETATION<br />
47<br />
willows in <strong>Yellowstone</strong><br />
National Park. Some are<br />
difficult to identify when<br />
specil11:ens are available for<br />
examination. Thus, historic<br />
photographs rarely allow for<br />
species identification, but do<br />
provide gross evidence of<br />
changes in abundance or<br />
distribution. Many observers<br />
have attributed the decline in<br />
willows on the northern range<br />
to overbrowsing by elk<br />
(Grimm 1939, Kay 1990,<br />
Chadde and Kay 1991, Wagner<br />
et al. 1995a). Other suggested causes include<br />
declines in beaver since the 19208 (Singer et al.<br />
1994), the colonization of the northern range by<br />
moose early in the 1900s (Houston 1982; Chadde<br />
and Kay 1988, 1991), fire suppression activities by<br />
park managers since the late 1800s (Houston 1973,<br />
1982), and climatic variations, especially the<br />
drought of the 19308, which, tlu-ougb changes in<br />
groundwater levels or other processes, might have<br />
either directly killed the plants or reduced their<br />
ability to produce secondary defensive chemical<br />
compounds that make them less palatable to<br />
grazers (Houston 1982; Singer et al. 1994, 1996b).<br />
Elk are usually the proximal factor in the<br />
decline of willow stands (Figure 4.3). Elk very<br />
likely accelerate willow declines that might be due<br />
to other causes. However, not all declines in<br />
willows are due to ungulate browsing. Houston<br />
(1982) pointed out that an entire stand of willows<br />
on tl,e lower northern range was defoliated by<br />
Disonycha pluriligata, a native beetle whose<br />
presence on the northern range was unknown until<br />
his study; he suggested that it would have been<br />
easy for the casual observer to mistake the aftereffects<br />
of this defoliation for heavy browsing by<br />
ungulates. In another instance, willow stands and<br />
individual willows died in the summers of 1988-<br />
1989 apparently as a consequence of the severe<br />
drought of 1988. Those dying stands were measured<br />
for water pressure in 1988 and had the lowest<br />
measurements of any of the many willow stands<br />
measured that year, verifying the drought stress<br />
(Singer et al. 1994).<br />
A third alternate factor in willow declines is<br />
fire. Willows on several marked transects burned<br />
and were either killed outright in 1988 or never resprouted<br />
or recovered (Norland et al. 1996). This<br />
is contrary to what others have observed. According<br />
to Wolff (1978) and MacCracken and Viereck<br />
(1990), willows virtually always vigorously resprout<br />
and reseed following fire. Some burned<br />
<strong>Yellowstone</strong> willow stands were immediately<br />
replaced with grasslands or forb vegetation. We<br />
interpret this as additional evidence that willows<br />
cannot recruit in many areas of the northern range<br />
and are failing on some sites under existing climate<br />
conditions or because of low surface/ground water<br />
control.<br />
Popular perceptions of willows, as with<br />
aspen (see next section), seem to be of a progressively<br />
declining group of species that has been<br />
fading from the northern range since early in this<br />
century. However, over the course of this century,<br />
most willow declines on the northern range<br />
occuned in the 1930s. Most ofthese occuned<br />
during the extended drought of that period, and in<br />
the face of elk numbers much lower than at present<br />
(Houston 1982, Singer 1996b). These declines<br />
included the disappearance of some willow stands<br />
and suppression of the heights of some other<br />
willow stands. Since then, there has been little<br />
change in willow status, perhaps lending credence<br />
to the idea that climatic forces, rather than ungulate<br />
browsing, may be the most important factor in<br />
Figure 4.3. Willows<br />
are protected from<br />
allllngll/ate<br />
brolVsing ill the<br />
Junction Butte<br />
exclosltre. Casual<br />
observation of this<br />
and other excloSllres<br />
has led to a mistaken<br />
impression that all<br />
vegetation all the<br />
northern range<br />
"should" have the<br />
same appearance as<br />
that inside<br />
exclosures. The only<br />
way the vegetation<br />
within the exclosllres<br />
could be replicated<br />
across the whole<br />
northern range<br />
would be by<br />
completely<br />
eliminating<br />
IInglllates from the<br />
range. NPS photo.