Forest-floor treatments in Arizona ponderosa pine restoration ecosystems: no short-term effects on plant communities

Leaf litter accumulation during fire exclusion and increases in tree density in postsettlement southwestern Pinus ponderosa forests may limit the establishment of understory vegetation. We performed an experiment in P. ponderosa forests of northern Arizona to ascertain plant community responses to forest-floor scarification and Oi removal on thirty-six 100-m 2 plots overlaid on an existing ecological restoration experiment that involved tree thinning and prescribed burning. Constrasting with findings from many other forest types, forest-floor treatments had no effect on community diversity or composition during the 2-year experiment. S?rensen similarities were as high as 97% between posttreatment years within treatments; and successional vectors also provided little indication that treatments may appreciably affect longer-term successional trajectories. Lack of response to these fairly drastic treatments is surprising given these forests' exceptionally heavy Oi horizons and large proportions of conifer litter. Apparently shading, belowground competition for water or nutrients, or other tree-associated factors more strongly limit understory communities than does leaf litter. Based on sparse A-horizon seed banks averaging <300 seeds ? m –2 and limited aboveground vegetation, we hypothesize that seed shortages, particularly for native perennials, also partly precluded a treatment response. Because extensive unvegetated areas at these restoration sites may be colonized by exotics, conservative management strategies could include testing the seeding or outplanting of desirable native species as an option for filling unoccupied microsites. Reporting of "no treatment effect" experiments such as this one is important to avoid biasing meta-analyses, as is future research to clarify combinations of factors limiting understory communities. Increased understanding of these limiting factors may lead to identification of other treatments that promote recovery of native species during ecological restoration in this region.     

Mycorrhization of ponderosa pine in a second-growth Sierra Nevada forest

Ectomycorrhizal colonization and rooting characteristics were quantified in a mature ponderosa pine ( Pinus ponderosa Dougl. ex Laws.) stand in the western Sierra Nevada. Root length totaled 3835.9 m · m –2 of forest floor surface area, with 96% consisting of the fine-root fraction. Total root dry weight and volume were 2230.4 g · m –2 and 5807.4 cm 3 · m –2 of forest floor area, respectively, with 69% of the former and 75% of the latter accounted for by the coarse fraction. Fine roots were most prevalent in the upper 15 cm of the mineral soil profile, and their abundance declined with increasing depth. Coarse roots were most abundant at a depth of 15–30 cm. Ectomycorrhizal counts totaled 26,814 · m –2 of forest floor area, and an overwhelming preponderance was associated with the fine-root fraction. More than three-quarters of mychorrhizae resided in the upper 15 cm of mineral soil, with an overall trend of declining numbers with increasing depth. Roots and mycorrhizae were exceedingly scarce at a depth of 45–60 cm, and neither was found in the organic soil layer above the mineral profile. A necessary step in understanding the ecophysiological role of mycorrhizae in mature forests is to quantify their abundance in such settings, and the results of this study contribute such information for ponderosa pine.     

Response of understory species to changes in ponderosa pine stocking levels in the Black Hills

The objective of this study was to test the hypothesis that there are no differences in understory production, by species, due to stocking levels of Pinus ponderosa (ponderosa pine). Understory production was estimated, by species, on 3 replicates each of 8 growing stock levels, ranging from clearcuts to unthinned stands, in both sapling- and pole-sized pine stands (48 plots) over 3 nonconsecutive years. All stands were approximately 70 yr old when thinning treatments were applied. Production of many herbaceous species, especially Agropyron spp. (wheatgrasses) and Carex spp. (sedges), declined as growing stock levels (measured in terms of basal area) of ponderosa pine increased. While trends in total production were similar, there were specific differences between sapling and pole stands. Sedges and Oryzopsis asperfolia (roughleaf ricegrass) produced more in sapling stands, whereas Danthonia intermedia (timber oatgrass) was more abundant in pole stands. Shrub production, dominated by Acrtostaphylos uva-ursi (bearberry), was relatively consistent across all stocking levels except unthinned. Although the total number of species declined as pine basal area increased, a few species, such as Linnaea borealis (twinflower) and Shepherdia canadensis (buffaloberry), were found only under relatively dense pine canopies. While floristic species richness was greater at lower stocking levels of ponderosa pine, the total number of species would be greater if all stocking levels were present.     

Wood and understory production under a range of ponderosa pine stocking levels, Black Hills, South Dakota

Stemwood and understory production (kg ha -1 ) were estimated during 3 nonconsecutive years on 5 growing stock levels of ponderosa pine including clearcuts and unthinned stands. Stemwood production was consistently greater at mid- and higher pine stocking levels, and understory production was greater in stands with less pine; however, there were no differences in total (stemwood + understory) production. Based on loss of productivity, there is no argument that small clearcuts and unthinned stands should not be included in site plans. They contribute significantly to community structure, particularly to plant and animal species richness.     

Production, losses, and germination of Ceanothus fendleri seeds in an Arizona ponderosa pine forest

I quantified seed production and ovule losses for Ceanothus fendleri Gray (Fendler ceanothus) plants protected from large ungulate herbivores in a ponderosa pine ( Pinus ponderosa [Laws]) forest of northern Arizona. I also tested seed germination responses to cold stratification and heat treatments in the laboratory. Fruit production on fecund stems ranged from 7.4 to 38.2 fruits per stem, which equated to 22.2-118.2 potential seeds based on 3 ovules per fruit. Stems that produced fruit were significantly large relative to their expected sizes. Predispersal ovule losses ranged from 70.7% to 91.4% across the 2 years studied. A chalcidoid seed parasite ( Eurytoma squamosa Bugbee) consumed 11%-28% of the total number of seeds produced. Postdispersal seed predation varied from 0% to 24% and was significantly affected by forest floor substrate in 1 study year. Cumulative ovule losses were estimated to be 71%-92%. Cold stratification did not significantly affect seed germination, and exposure to 90°C resulted in the highest germination percentage. Both dormant and nondormant seeds suggested a bet-hedging life history strategy. This study provides basic ecological information important for management of ponderosa pine forest and nursery production of C. fendleri .     

A seed chalcid ( Eurytoma squamosa Bugbee) parasitizes buckbrush ( Ceanothus fendleri Gray) seeds in a ponderosa pine forest of Arizona

Predispersal seed parasitism rates were quantified for buckbrush ( Ceanothus fendleri Gray) in 3 Arizona ponderosa pine ( Pinus ponderosa Laws.) forest units that had been thinned for ecological restoration objectives. The chalcidoid wasp Eurytoma squamosa Bugbee (Eurytomidae) was responsible for 35% of total seed loss in a single year. These findings represent an expansion of the known range and host list for E. squamosa and increase our understanding of factors that may constrain regeneration of C. fendleri in Southwest ponderosa pine forests.     

Natural variability of vegetation, soils, and physiography in the bristlecone pine forest of the Rocky Mountains

Pinus aristata Engelm. forest vegetation in Colorado was studied to determine vegetation composition and the relationship between vegetation and environment. Species percent cover, tree size class, and environmental variables were recorded for 49 plots. Previously collected data for 4 plots form New Mexico were included. Environmental variables included latitude, longitude, elevation, slope, aspect, topographic position, parent material, percent rock cover, mean rock size, litter depth, estimated plot age class, and evidence of anthropogenic disturbance. Soils were analyzed for texture, depth, and percent carbon and nitrogen. Pinus aristata foliage was analyzed for percent nitrogen and phosphorus. Direct and indirect gradient analyses (CANOCO) were used to determine environmental factors associated with community composition. Vegetation in P. aristata forests is influenced primarily by elevation and soil pH. Substrate, soil texture, topographic position, and geographic location are secondary factors. Six plant associations were identified using cluster analysis (listed in an elevational sequence from low to high): Pinus aristata/ Festuca arizonica Vasey, Pinus aristata / Festuca thurberi Vasey, Pinus aristata / Juniperus communis L., Pinus aristata / Vaccinium myrtilus L., Pinus aristata / Ribes montigenum McClatchie, and Pinus aristata / Trifolium dasyphyllum Torr. & Gray.