Mediterranean climate effects. II. Conifer growth phenology across a Sierra Nevada ecotone

Growth and xylem water potential of the lower elevation conifers Pinus jeffreyi and Abies concolor and the higher elevation Pinus monticola and Abies magnifica were monitored in their montane Mediterranean habitat of the southernmost Sierra Nevada mountains of California. Measurements were made across the ecotone between the midmontane and upper montane forests and through light and heavy snowfall years.Radial stem growth, averaging ～1.5 mm/yr, started 2 wk after snow melt, providing that maximum air temperatures had reached 21°C, and ended when predawn water potentials fell rapidly at the onset of the summer drought. Leader growth started on or after a fixed date, providing that minimum air temperatures were above -4°C for Pinus species or +2.5°C for Abies species. The cue for leader growth was inferred to be photoperiodic. Leader growth ended when either a determinate internode length of ～1 mm was reached or predawn water potentials fell rapidly. Abies magnifica grew more rapidly than the low-elevation species, but had a shorter growth period; its annual leader growth, as a consequence, was only 35 mm/yr vs. 50 mm/yr for the low-elevation species. Needle growth was similarly determinate in the absence of early drought. This growth phenology contributes to determining species distribution across the ecotone.     

Water source utilization by Pinus jeffreyi and Arctostaphylos patula on thin soils over bedrock

Stable isotopes were used to evaluate water sources for co-occurring Jeffrey pine (Pinus jeffreyi Grev & Balf.) and greenleaf manzanita (Arctostaphylos patula Greene) in the southern Sierra Nevada, California, where soils averaged only 75 cm thick but were underlain by up to 5 m of weathered granitic bedrock. Soils and underlying weathered bedrock were sampled three times during both the 1997 and 1998 growing seasons, in 25 cm increments, from 0 to 400 cm or until hard bedrock was reached, and plant stem tissue was sampled simultaneously. Extracted water from the soil/bedrock substrate and plant tissue was analyzed for delta(18)O and/or deltaD, and depth of water source was determined by inference in conjunction with moisture status of the substrate. Water source utilization over the growing seasons for both plants generally followed a pattern similar to that observed for water depletion. Predominant water use was initially from the surface soils. Progressively deeper water sources, including weathered bedrock to a depth of several meters, were exploited as the season progressed and the overlying substrate was depleted of moisture. Early in the growing season, stable isotope values were slightly lower for pine than for manzanita (e.g., average deltaD in June 1997 was -81 per thousand for pine and -77 per thousand for manzanita), and suggest that the functional rooting depth for pine may have been slightly greater than for manzanita. In September 1997, manzanita deltaD values averaged -57 per thousand while pine values averaged -85 per thousand, indicating that manzanita opportunistically utilized summer precipitation while pine used more dependable bedrock water. In 1998, soils remained moist through July due to a late snowfall. Unlike the previous year, pine and manzanita deltaD values were not significantly different in mid- and late-growing season, and both plants exploited bedrock-derived water as soil water was depleted. Water held within bedrock was essential for meeting plant transpirational requirements over the summer drought.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Water use patterns of four co-occurring chaparral shrubs

Summary Mixed stands of chaparral in California usually contain several species of shrubs growing close to each other so that aerial branches and subterranean roots overlap. There is some evidence that roots are stratified relative to depth. It may be that root stratification promotes sharing of soil moisture resources. We examined this possibility by comparing seasonal water use patterns in a mixed stand of chaparral dominated by four species of shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a neutron probe and soil phychrometers to follow seasonal depletion and recharging of soil moisture and compared these patterns to seasonal patterns of predawn water potentials, diurnal leaf conductances, and diurnal leaf water potentials. Our results indicated that 1) Quercus was deeply rooted, having high water potentials and high leaf conductances throughout the summer drought period, 2) Heteromeles/Adenostoma were intermediate in rooting depth, water potentials, and leaf conductances, and 3) Rhamnus was shallow rooted, having the lowest water potentials and leaf conductances. During the peak of the drought, predawn water potentials for Quercus corresponded to soil water potentials at or below a depth of 2 m, predawn water potentials of Heteromeles/ Adenostoma corresponded to a depth of 0.75 m, and predawn water potentials of Rhamnus corresponded to a depth of 0.5 m. This study supports the concept that co-occurring shrubs of chaparral in California utilize a different base of soil moisture resources.     

Forest/environment relationships in Yosemite National Park,California, USA

Forest compositional patterns in Yosemite National Park, California, were related to environmental factors through numerical classification of forest types, arrangement of forest types along elevational and topographic gradients, and development of regression models relating basal area of common tree species to environmental variables. The eight forest types are differentiated primarily by elevation zone and secondarily by topographic setting. Lower montane forests (1200–1900 m) were divided into the Abies concolor/Calocedrus type occurring primarily on mesic sites and the Pinus ponderosa/Calocedrus type predominantly on xeric sites. Upper montane forests (1900–2500 m) included the Abies concolor/Abies magnifica type on mesic sites, the Abies magnifica/Pinus type on somewhat more xeric sites, and Juniperus occidentalis/Pinus jeffreyi woodlands on granitic domes. Subalpine forests (2500–3300 m) embraced three types: Tsuga mertensiana/Pinus forests on mesic sites, monotypic Pinus contorta forests on drier sites, and Pinus albicaulis/Pinus contorta groves at treeline. Regression models consistently included elevation and soil magnesium content as explanatory variables of species basal area totals. The two Abies spp. were negatively correlated with soil magnesium levels, whereas other montane species (e.g. Calocedrus decurrens, Pinus lambertiana, and Pinus ponderosa) exhibited positive correlation with soil magnesium. Topography and soil physical properties were only infrequently incorporated into species regression models.Abbreviations DBH= diameter at breast height (1.4 m) - DCA= detrended correspondence analysis - TWINSPAN= two-way indicator species analysis     

SURVIVAL OF SEVERE DROUGHT BY A NON-SPROUTING CHAPARRAL SHRUB

A 2-yr drought (1975-1977) in much of California provided unusually severe water stress on many native plant species. In this paper we evaluate the effects of this drought on Arctostaphylos viscida Parry, a common non-sprouting chaparral shrub in the foothills of the southern Sierra Nevada. At the peak of the drought, water potentials as low as –74 bars were measured. While only three of 90 shrubs sampled were killed by the drought, all but one showed signs of drought-induced dieback. First and second year post-drought twig growth was significantly greater on shrubs with 90% or greater branch mortality than on those suffering lesser dieback. In all cases, new growth occurred only on surviving branches. Midday water stress measurements showed little difference for shrubs exhibiting high and low levels of dieback. These results suggest that drought survival in Arctostaphylos viscida may depend on a mechanism where certain branches are sacrificed with the surviving ones preferentially receiving available resources. The possibility that this may involve the dieback of below ground roots and/or lack of production of new roots to compensate for lack of root turnover is supported by water stress measurements of adjacent fire damaged shrubs. Implications of possible specialized drought survival mechanisms in non-sprouting shrubs are explored.     

Effects of plant size on photosynthesis and water relations in the desert shrub Prosopis glandulosa (Fabaceae)

The Jornada del Muerto basin of the Chihuahuan Desert of southern New Mexico, USA, has undergone a marked transition of plant communities. Shrubs such as mesquite (Prosopis glandulosa) have greatly increased or now dominate in areas that were previously dominated by perennial grasses. The replacement of grasses by shrubs requires an establishment phase where small shrubs must compete directly with similar-sized grass plants. This is followed by a phase in which large, established shrubs sequester nutrients and water within their biomass and alter soil resources directly under their canopy, creating “islands” of fertility. We hypothesized that these two phases were associated with shrubs having different physiological response capacities related to their age or size and the resource structure of the environment. As a corollary, we hypothesized that responses of small shrubs would be more tightly coupled to variation in soil moisture availability compared to large shrubs. To test these hypotheses, we studied gas exchange and water relations of small (establishing) and large (established) shrubs growing in the Jornada del Muerto as a function of varying soil moisture during the season. The small shrubs had greater net assimilation, stomatal conductance, transpiration, and xylem water potential than large shrubs following high summer rainfall in July, and highest seasonal soil moisture at 0.3 m. High rates of carbon assimilation and water use would be an advantage for small shrubs competing with grasses when shallow soil moisture was plentiful. Large shrubs had greater net assimilation and water-use efficiency, and lower xylem water potential than small shrubs following a dry period in September, when soil moisture at 0.3 m was lowest. Low xylem water potentials and high water-use efficiency would allow large shrubs to continue acquiring and conserving water as soil moisture is depleted. Although the study provides evidence of differences in physiological responses of different-sized shrubs, there was not support for the hypothesis that small shrubs are more closely coupled to variation in soil moisture availability than large shrubs. Small shrubs may actually be less coupled to soil moisture than large shrubs, and thus avoid conditions when continued transpiration could not be matched by equivalent water uptake.     

毛乌素沙地南缘沙柳灌丛土壤水分及水量平衡

以毛乌素沙地南缘沙柳人工固沙灌丛为研究对象,对不同栽植密度(0.2、0.6和0.8株·m-2)沙柳灌丛生长季土壤水分动态和蒸散量变化进行研究.结果表明:不同栽植密度沙柳灌丛区土壤水分动态和蒸散量存在明显差异,土壤含水量随着栽植密度增加呈单峰型曲线;生长季内沙柳灌丛土壤含水量变化呈“S”形曲线,并与降雨存在密切的关系.蒸散量以栽植密度0.8株·m-2的沙柳灌丛最高(114.5 mm),占同期降雨量的90.8％;以0.6株·m-2的沙柳灌丛最低(109.7 mm).根据生长季土壤水分动态和水分平衡特征,毛乌素沙地南缘沙柳灌丛适种密度为0.6株·m-2.     

科尔沁沙地不同地形小叶锦鸡儿灌丛土壤水分动态

根据科尔沁沙地广泛分布的小叶锦鸡儿灌丛的植被调查和土壤水分监测数据,分析了沙地人工固沙灌丛的土壤水分时空动态和土壤储水量变化,并应用水量平衡法测定了灌丛蒸散发.结果表明:丘间地的土壤水分条件最好,丘中次之,丘上最低;灌丛区的土壤水分含量随深度增加而增加,在生长季内不会发生水分胁迫.灌丛区土壤水分与降雨过程高度相关,深层(50～180 cm)土壤水分同降雨的相关性高于表层(0～50 cm)土壤,并且深层土壤水分的变异也大于表层.整个生长期内,小叶锦鸡儿灌丛土壤储水量增加,土壤水分处于积累中,估算蒸散量占同期降雨量的64%以上.     

Effects of young Artemisia rothrockii shrubs on soil moisture,soil nitrogen cycling,and resident herbs

Questions: How do young sagebrush shrubs (Artemisia rothrockii, Asteraceae) affect soil moisture availability? How do young sagebrush shrubs affect soil nitrogen cycling? How does the resident herb community respond to shrub removal in the early stages of sagebrush encroachment? Location: Mulkey and Bullfrog Meadows on the Kern Plateau in the Golden Trout Wilderness, Sierra Nevada Mountains, Inyo National Forest, Inyo County, California, USA. Methods: We removed young encroaching sagebrush shrubs from 3.5 m × 3.5 m plots and compared soil moisture, net mineralization, net nitrification, and herb cover with paired control plots over four growing seasons. Results: On average throughout the experiment, the difference between removal plots and control plots in soil moisture was small. Removal plots were wetter by 1.3 ± 2.0% at 0–30 cm depth, 2.1 ± 3.1% at 30–60 cm depth and 3.1 ± 5.8% at 60–90 cm depth. By contrast, after four years, net mineralization was 32 ± 26% (mean ± 95% CI) lower in sagebrush removal plots, suggesting that sagebrush encroachment increases rates of N‐cycling. Total herb cover was 13.0 ± 6.4% (mean ± 95% CI) higher in plots where young sagebrush shrubs were removed. This difference in cover appeared during the first season in which sagebrush shrubs were removed. Conclusions: Our results suggest that while young sagebrush shrubs do not contribute substantially to meadow drying, they alter N cycling rates, and may indirectly increase the rate of their own encroachment by competitively reducing resident herbs.     

Influence of soil thickness on stand characteristics in a Sierra Nevada mixed-conifer forest

Soil thickness can be an important factor influencing vegetation, yet few spatially-explicit studies have examined soil horizon thickness and vegetation composition in summer-drought forests. We compared seismic and soil penetration measurements of combined A + C and Cr horizon thickness, soil moisture and temperature, and stand variables in a contiguous 4-ha mixed-conifer stand of the Sierra Nevada. Thickness of A + C and Cr horizons were highly variable but were not correlated to each other. Total basal area and canopy cover were positively related with A + C horizon thickness, and shrub cover was positively related with Cr horizon thickness. Basal area of white fir [Abies concolor (Gord and Glend) Lindl.] and incense-cedar [Calocedrus decurrens (Torrey) Florin] were positively correlated with A + C horizon thickness, but there was no relationship between A + C or Cr horizon thickness and basal area of Jeffrey pine (Pinus jeffreyi Grev. and Balf.), sugar pine (P. lambertiana Douglas), or red fir (A. magnifica A. Murray). Both white and red fir seedlings were associated with decreased soil temperature, but only white fir seedlings were positively associated with soil moisture. Soil penetration estimates of soil thickness were similar to seismic estimates for shallow soils (<50 cm depth) but were poorly related on deeper soils. Visual surface conditions and tile probe estimates of soil thickness can be highly misleading because ‘shallow’ areas may have a thick layer of weathered bedrock that can serve as a potential rooting medium for deep-rooted trees and shrubs. In our study only the refraction seismic method had the potential to measure total soil depth that included A + C and Cr horizon thickness.     

Gradient Analysis of Fire Regimes in Montane Forests of the Southern Cascade Range,Thousand Lakes Wilderness,California, USA

Species distribution and abundance patterns in the southern Cascades are influenced by both environmental gradients and fire regimes. Little is known about fire regimes and variation in fire regimes may not be independent of environmental gradients or vegetation patterns. In this study, we analyze variation in fire regime parameters (i.e., return interval, season, size, severity, and rotation period) with respect to forest composition, elevation, and potential soil moisture in a 2042 ha area of montane forest in the southern Cascades in the Thousand Lakes Wilderness (TLW). Fire regime parameters varied with forest composition, elevation, and potential soil moisture. Median composite and point fire return intervals were shorter (4-9 yr, 14-24 yr) in low elevation and more xeric white fir (Abies concolor)-sugar pine (Pinus lambertiana) and white fir-Jeffrey pine (P. jeffreyi) and longest (20-37 yr, 20-47 yr) in mesic high elevation lodgepole pine (Pinus contorta) and red fir (Abies magnifica)-mountain hemlock (Tsuga mertensiana) forests. Values for mid-elevation red fir-white fir forests were intermediate. The pattern for fire rotation lengths across gradients was the same as for fire return intervals. The percentage of fires that occurred during the growing season was inversely related to elevation and potential soil moisture. Mean fire sizes were larger in lodgepole pine forests (405 ha) than in other forest groups (103-151 ha). In contrast to other parameters, fire severity did not vary across environmental and compositional gradients and >50% of all forests burned at high severity with most of the remainder burning at moderate severity. Since 1905, fire regimes have become similar at all gradient positions because of a policy of suppressing fire and fire regime modification will lead to shifts in landscape scale vegetation patterns.     

MATING SYSTEM IN NATURAL POPULATIONS OF JEFFREY PINE

The mating system and allozyme variation at 20 loci in three Klamath Mountains and two Sierra Nevada populations of Jeffrey pine (Pinus jeffreyi Grev. & Balf.) were investigated. On average, multilocus estimates of the proportion of viable progeny due to outcrossing (t_m) were high in all populations (mean t_m = 0.935, range 0.881 to 0.971). Despite differences in stand structure, t_m did not differ (P > 0.05) between the Klamath (mean t_m = 0.933) and Sierra Nevada (mean t_m = 0.937) populations. At all but one locus in one population and at two in another, genotype frequencies fit (P > 0.05) Hardy-Weinberg expectations. Mean estimates of observed heterozygosity in Klamath (0.182) and Sierra Nevada (0.327) populations were comparable to values reported for other conifers.     

Nesting habitat of Warbling Vireos across an elevational gradient in the southern Sierra Nevada

ABSTRACT.   Populations of Warbling Vireos ( Vireo gilvus ) are declining in California, apparently due to low reproductive success. From 1989–2002, I studied the nest-site selection and reproductive success of Warbling Vireos across an elevational gradient in the southern Sierra Nevada. Warbling Vireos regularly nested in upland coniferous forests with few or no deciduous trees, and tree species used by nesting vireos included five species of conifers and four species of deciduous trees. Overall, hardwoods were used more than expected based on their availability, but 69% of all nests were in conifers. Hardwood trees were found only in low and mid-elevation ponderosa pine ( Pinus ponderosa ) and mixed-conifer sites. In low-elevation ponderosa pine habitat, 87% of nests were in hardwoods, with 67% in California black oaks ( Quercus kelloggii ), a species that typically occupies upland sites. In mixed-conifer sites where reproductive success was high, 65% of nests were in incense cedar ( Calocedrus decurrens ) and California black oak was the next most commonly used species. Because fire suppression has likely increased numbers of shade-tolerant tree species like incense cedar, shade-intolerant species like black oaks may have been more important as a nest substrate for vireos in the past. Only conifers were used as nesting substrates at higher elevations. Nest success was greater for Warbling Vireos that nested in tall trees in areas with high basal area. My results suggest that Warbling Vireos in the Sierra Nevada would benefit from management activities that encourage retention and recruitment of California black oaks at lower elevations, and development of stands with large trees, dense foliage, and semi-open canopy throughout their elevation range.     

Benefits of Using Shrubs as Nurse Plants for Reforestation in Mediterranean Mountains: A 4-Year Study

Shrubs are commonly considered competitors of planted seedlings in reforestation programs. However, shrubs can facilitate the establishment of understory seedlings in environments that, like Mediterranean‐type ecosystems, are characterized by harsh environmental conditions. In 1997, an experiment was set up in the Sierra Nevada mountains (southeast Spain) to test the use of shrubs as nurse plants for an alternative reforestation technique. Two‐year‐old seedlings of Pinus sylvestris and Pinus nigra were planted in four microhabitats: (1) open interspaces without vegetation (which is the usual method employed in reforestation programs), (2) under individuals of the shrub Salvia lavandulifolia, (3) under the north side of spiny shrubs, and (4) under the south side of spiny shrubs. Seedlings were also distributed in plots with and without ungulates to test the effect of herbivore damage. We report here the results of survival and growth after four growing seasons, a time span long enough to draw robust conclusions concerning the suitability of this technique. Pine survival was remarkably higher when planted under individuals of S. lavandulifolia as compared with open areas (2.6 times for P. sylvestris and 1.8 for P. nigra). The survival of both pine species was also higher when planted on the north side of spiny shrubs, while mortality on the south side was similar to that found in open areas. The reduction of solar radiation by the canopy of shrubs was likely the main factor determining shrub facilitation. The growth of the pines differed among years. However, growth was not inhibited when planted with shrubs as compared with open areas in any of the years. Herbivore damage was low but was mostly concentrated in the leader shoot, exacerbating the deleterious effect of ungulate herbivores on pine growth. We conclude that the use of shrubs as nurse plants for reforestation is a viable technique to increase establishment success of reforestation in Mediterranean‐type ecosystems and that it might be similarly useful in other water‐stressed environments. In addition, this technique offers the advantage of following natural succession, thus minimizing the impact in the community.     

Depth of water acquisition by invading shrubs and resident herbs in a Sierra Nevada meadow

Woody plant encroachment into semiarid ecosystems has become a global trend in recent decades. Due to stream channel incision, the semiarid riparian montane meadows of the southern Sierra Nevada Mountains, USA are experiencing long-term declines in soil moisture. A woody shrub, Artemisia rothrockii A. Gray (Rothrock sagebrush, Asteraceae) is invading these herbaceous meadows. We used an analysis of the stable oxygen isotope ratios of plant and soil water to measure the depth of plant water acquisition during the early stages of this woody plant encroachment. Sagebrush used deeper water on average than most herbs, but it also acquired 10–30% of its water from shallow (<30 cm) soil. Most of the young sagebrush seedlings (1–3 years old, <15 cm) that we sampled used deep water like the older shrubs. Many, but not all of the herb species we sampled were also able to acquire deep water. These findings are consistent with a scenario of shrub encroachment in which channel incision causes shallow-water-dependent herbs to die back, allowing shrub seedlings to establish in disturbed areas during wet years. At least during the early stages of the invasion, some herbs appear to coexist with sagebrush by using deep root systems to cope with the declining shallow soil moisture.     

A unique ballistosporic hypogeous sequestrate Lactarius from California

Lactarius rubriviridis sp. nov., characterized by hypogeous, sequestrate basidiomes with red latex, green stains, and forcibly discharged, reticulate basidiospores is described and illustrated. During the Spring, the new species forms basidiomes associated with conifers at 1400-1800 m elevation in the Sierra Nevada, and is known from two specimens collected 19 yr apart. Comparisons with the putatively polyphyletic genera Arcangeliella and Zelleromyces, and an accounting of all known members of these genera are provided.     

Interspecific variation in sapling mortality in relation to growth and soil moisture

To examine the causes of landscape variation in forest community composition, we have quantified sapling mortality as a function of growth and soil moisture for seven dominant species in transition oak-northern hardwood forests of the northeastern USA. We located saplings in sites that encompassed a wide range of variation in soil moisture and light availability. In mesic conditions, the probability of mortality decays rapidly with increasing growth among shade tolerant species and more gradually among shade intolerant species: the rank order of survivorship at low growth rates is Tsuga canadensis > Fagus grandifolia > Acer saccharum > Fraxinus americana > Acer rubrum > Quercus rubra > Pinus strobus . The relationship between probability of mortality and growth does not vary with soil moisture among species insensitive to drought: Tsuga canadensis , Quercus rubra, and Pinus strobus . However, probability of mortality increases substantially with decreasing soil water availability for the other four species. Acer saccharum and Fagus grandifolia have high mortality rates under xeric conditions even when their growth is not suppressed. Acer rubrum and Fraxinus americana exhibited a steady but more gradual increase in the probability of mortality with decreasing soil moisture. Among the five deciduous hardwood species we examined there is a weak inverse relationship between the ability to survive growth suppression, a measure of shade tolerance, and the ability to survive in xeric conditions, a measure of drought tolerance. Tsuga canadensis , however, is tolerant of growth suppression and exhibits high survivorship in xeric conditions, while Pinus strobus is intolerant of growth suppression but insensitive to soil moisture. Species differences in water-dependent mortality are consistent with the species distributions across landscape gradients of soil water availability.     

Canopy vs. soil effects of shrubs facilitating tree seedlings in Mediterranean montane ecosystems

Abstract. Question: Is the facilitative effect of nurse shrubs on early recruitment of trees mediated by a ‘canopy effect’(microclimate amelioration and protection from herbivores), a ‘soil effect’(modification of soil properties), or both? Location: Two successional montane shrublands at the Sierra Nevada Protected Area, SE Spain. Method: Seedlings of Quercus and Pinus species were planted in four experimental treatments: (1) under shrubs; (2) in open interspaces without vegetation; (3) under shrubs where the canopies were removed; (4) in open interspaces but covering seedlings with branches, mimicking a shrub canopy. Results: Both effects benefited seedling performance. However, microclimatic amelioration due to canopy shading had the strongest effect, which was particularly pronounced in the drier site. Below‐ground, shrubs did not modify soil physical characteristics, organic matter, total N and P, or water content, but significantly increased available K, which has been shown to improve seedling water‐use efficiency under drought conditions. Conclusions: We propose that in Mediterranean montane ecosystems, characterised by a severe summer drought, pioneer shrubs represent a major safe site for tree early recruitment during secondary succession, improving seedling survival during summer by the modification of both the above‐and below‐ground environment.     

Conifer colonization of a 350-year old rock fall at Lassen Volcanic National Park in northern California

Montane conifers of western North America commonly colonize mineral surfaces resulting from landslides, lahars, mudflows, and rock avalanches. This colonization can include shade-tolerant conifers that may eventually dominate the forest in a pattern termed “direct” succession. Documenting examples of this long-term successional process are useful for identifying alternative successional trajectories and indicating potential controlling mechanisms for subsequent experimental analysis. This study (1) analyzes the 1992 status of the conifer colonization on the coarse-textured surface of a 1650 AD rock avalanche in northern California and (2) measures individual growth and survivorship in permanent plots between 1992 and 2003. Increment cores of large trees indicated initial conifer colonization before 1700 AD with continuous subsequent colonization. Mean conifer density in 1992 was 725 (SD = 747) ha⁻¹ with densities increasing with decreasing rock sizes. Densities were not correlated with distances to possible seed sources. Median heights were <1 m, and the mean proportional height growth rate for healthy individuals was 0.0166 year⁻¹. The mortality rate for individuals ≥0.1-m tall was <0.007 year⁻¹. The conifers were more numerous than shrubs, and there was little apparent evidence of facilitation or inhibition of conifers. The species assemblage is mostly (89%) Abies concolor, Pinus monticola, and Pinus ponderosa individuals dispersed across an elevation range of 1870–2040 m a. s. l. This is an atypical species mix for these elevations in this location, and this mix is not readily predicted from species properties such as seed mass, seed flight distance, or drought tolerance.