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Yellowstone's Northern Range - Greater Yellowstone Science ...

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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.

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