Forest vegetation of the Colorado Front Range

Abstract

The forest vegetation of the northern Colorado Front Range was studied using a combination of gradient analysis and classification methods. A graphical model of forest composition based on elevation and topographic-moisture gradients was constructed using 305 0.1 ha samples. To derive the topographic moisture gradient, stands were stratified into eight 200 m elevation belts, and then ordinated by correspondence analysis using understory (<1 m) data. Each of the resultant gradients was scaled against a standard site moisture scalar derived from incident solar radiation and topographic position. Except for krummholz sites, the vegetation defined gradients fit the moisture scalar closely. Once scaled, these gradients were stacked vertically, sandwich-style, to create the graphical representation shown in Figure 5.

Gradient analysis and ordination (direct and indirect gradient analysis of Whittaker, 1967) are frequently viewed as alternative approaches for analysis of vegetation. With gradient analysis the axes are readily interpretable, but stand placement is often difficult and at times questionable. Ordination defines an optimal arrangement for species and/or stands, but axis interpretation is often impossible. With the present combination of methods, the interpretability of gradient analysis complements the precision of placement obtained with ordination.

Forest vegetation was classified by dividing the gradient model into eight series and 29 types on the basis of similar successional trends in canopy dominants. On dry, low-elevation sites above 1 700 m Pinus ponderosa woodlands dominate. With increasing elevation or site moisture, tree density increases and Pinus ponderosa, Pseudotsuga forests prevail. At middle elevations on mesic sites forests of mixed composition occur. Pinus contorta forests dominate at middle elevations over much of the central position of the moisture gradient, though these are primarily post-fire forests. With protection from fire only a small percentage of sites retain dominance by Pinus contorta. Over the lower portion of its range Pinus contorta is succeeded by Pseudotsuga, while at higher elevations Abies lasiocarpa and Picea engelmannii can eventually achieve dominance. At high elevations on all except the driest sites Picea engelmannii and Abies lasiocarpa are exclusive dominants, both after disturbance and in climax forests. Pinus flexilis dominates on the driest high-elevation sites. Above 3 500 m forests are replaced by alpine tundra, often with a transitional krummholz zone.

Structure and post-fire development were examined in the context of the gradient-based classification scheme. Three generalized types of forest development were recognized as reference points in a continuum of developmental patterns varying with both elevation and soil moisture.

On favorable, middle-elevation sites, trees become established rapidly after disturbance. Rapid growth results in severe overcrowding and competitive elimination of reproduction. As a consequence bell-shaped diameter distributions develop. Diversity and productivity appear to drop while biomass remains roughly constant. Following decades or even centuries of stagnation, the forests eventually breakup through mortality of the canopy trees, thereby allowing regeneration to resume. During this period of renewed regeneration, biomass, diversity, and productivity all show dramatic changes in response to the changing population structure (Fig. 9). This type of forest development can be found in forests dominated by Picea engelmannii and Abies lasiocarpa, Pinus contorta, Pseudotsuga menzeisii, Pinus flexilis or Populus tremuloides. On highest elevation forest sites or at middle elevations on the very driest sites reestablishment rates are greatly reduced. These forests dominated by Picea and Abies or Pinus flexilis gradually approach predisturbance levels of biomass, diversity and productivity, while regeneration remains at a roughly constant level. At lower elevations in the Pinus ponderosa woodlands, regeneration appears episodic, reflecting variation in seed rain and favorable conditions for seedling growth. Here, inter-tree competition is relatively unimportant and diameter distributions show irregular humps resulting from periodic recruitment.

A few species pairs presented consistent problems and their treatment as single species was necessary. Garex rossii and C. brevipes were lumped as Carex rossii. Rosa woodsii and R. acicularis were lumped as Rosa sp. Cirsium scopulorum and C. coloradense were lumped as Cirsium coloradense. Extreme forms of Arnica cordifolia and A. latifolia are easily distinguishable, but as these species intergrade and hybridize extensively, they have been lumped as Arnica cordifolia. The native bluegrass, Poa agassizensis, was lumped with Poa paratensis. Solidago missouriensis includes some S. canadensis.