Understory vegetation indicates historic fire regimes in ponderosa pine‐dominated ecosystems in the Colorado Front Range

Question: Can current understory vegetation composition across an elevation gradient of Pinus ponderosa‐dominated forests be used to identify areas that, prior to 20th century fire suppression, were characterized by different fire frequencies and severities (i.e., historic fire regimes)? Location: P. ponderosa‐dominated forests in the montane zone of the northern Colorado Front Range, Boulder and Larimer Counties, Colorado, USA. Methods: Understory species composition and stand characteristics were sampled at 43 sites with previously determined fire histories. Indicator species analyses and indirect ordination were used to determine: (1) if stands within a particular historic fire regime had similar understory compositions, and (2) if understory vegetation was associated with the same environmental gradients that influence fire regime. Classification and regression tree analysis was used to ascertain which species could predict fire regimes. Results: Indicator species analysis identified 34 understory species as significant indicators of three distinct historic fire regimes along an elevation gradient from low‐ to high‐elevation P. ponderosa forests. A predictive model derived from a classification tree identified five species as reliable predictors of fire regime. Conclusions: P. ponderosa‐dominated forests shaped by three distinct historic fire regimes have significantly different floristic composition, and current understory compositions can be used as reliable indicators of historical differences in past fire frequency and severity. The feasibility demonstrated in the current study using current understory vegetation properties to detect different historic fire regimes, should be examined in other fire‐prone forest ecosystems.     

Vegetation–environment relationships and ecological species groups of an Arizona Pinus ponderosa landscape, USA

Pinus ponderosa forests occupy numerous topographic and soil complexes across vast areas of the southwestern United States, yet few data exist on species distributions and vegetation–environment relationships for these environmentally diverse landscapes. We measured topography, soils, and vegetation on 66, 0.05-ha plots within a 110,000-ha P. ponderosa landscape in northern Arizona, USA, to discern vegetation–environment relationships on this landscape. We analyzed associations of environmental variables with plant communities and with single-species distributions, and we classified ecological species groups (co-occurring plant species exhibiting similar environmental affinities). Gradients in community composition paralleled gradients in soil texture, available water, organic C, total N, and geographic precipitation patterns. Soil parent material, affected by the presence or absence of volcanic activity, is a primary factor constraining vegetation patterns on this landscape. Using discriminant analysis, we built a model that correctly classified the most important of four grasses (Bouteloua gracilis, Muhlenbergia montana, Sporobolus interruptus, or Festuca arizonica) on 70–80% of plots based on five environmental variables related to soil moisture and resource levels. We also classified 52 of the 271 detected plant species into 18 ecological species groups. Species groups ranged from Phacelia and Bahia groups occupying xeric, volcanic cinder soils low in organic C and total N, to Festuca and Lathyrus groups characterizing moist, loam and silt loam soils. We applied the species groups by estimating P. ponderosa diameter increment in a regression tree using abundances of species groups. The most rapid P. ponderosa diameter growth of 5 mm/year occurred on plots with high importance of the Festuca and Lathyrus groups. Our results on this semi-arid landscape support several general ecological species group principles chiefly developed in temperate regions, and suggest that vegetation–environment research has great potential for enhancing our understanding of P. ponderosa forests occupying vast areas of the southwestern United States.     

Ethylene production by different age class ponderosa and Jeffery pine needles as related to ozone exposure and visible injury

Summary Ethylene produced by different needle age classes representing natural populations of two ponderosa pine varieties [Pinus ponderosa var. arizonica (Engelm.) Shaw and var. ponderosa Dougl. ex Laws.] and Jeffery pine (Pinus jeffreyi Grev. and Balf.) was characterized using mercuric perchlorate traps. All populations contained individual trees which were either symptomatic or asymptomatic with respect to visible ozone injury. Ethylene production by different needle age classes was also characterized in P. ponderosa var. ponderosa seedlings grown in open top ozone fumigation chambers. Older age class needles produce significantly (P>0.05) more ethylene than younger age class needles. Needles of both P. ponderosa var. ponderosa and P. jeffreyi exhibiting ozone injury in the field produced significantly (P>0.05) higher levels of ethylene than asymptomatic conspecific trees. Seedlings exposed to the highest level of ozone in the fumigation study produced the highest levels of ethylene, followed by fumigation with medium and low ozone concentrations and carbon filtered air. These data indicate that the measurement of ethylene in conifer needles, as a measure of stress, needs to be calibrated for needle age class. It also suggests that the sensitivity of a tree to ozone injury may be regulated by the inherent ability of the individual to produce ethylene.     

Forest ecosystems of an Arizona Pinus ponderosa landscape: multifactor classification and implications for ecological restoration

Aim We developed an ecosystem classification within a 110,000‐ha Arizona Pinus ponderosa P. & C. Lawson (ponderosa pine) landscape to support ecological restoration of these forests. Specific objectives included identifying key environmental variables constraining ecosystem distribution and comparing plant species composition, richness and tree growth among ecosystems. Location The Coconino National Forest and the Northern Arizona University Centennial Forest, in northern Arizona, USA. Methods We sampled geomorphology, soils and vegetation on 66 0.05‐ha plots in open stands containing trees of pre‐settlement (c. 1875) origin, and on 26 plots in dense post‐settlement stands. Using cluster analysis and ordination of vegetation and environment matrices, we classified plots into ecosystem types internally similar in environmental and vegetational characteristics. Results We identified 10 ecosystem types, ranging from dry, black cinders/Phacelia ecosystems to moist aspen/Lathyrus ecosystems. Texture, organic carbon and other soil properties reflecting the effects of parent materials structured ecosystem distribution across the landscape, and geomorphology was locally important. Plant species composition was ecosystem‐specific, with C₃Festuca arizonica Vasey (Arizona fescue), for instance, abundant in mesic basalt/Festuca ecosystems. Mean P. ponderosa diameter increments ranged from 2.3–4.3 mm year^?1 across ecosystems in stands of pre‐settlement origin, and the ecosystem classification was robust in dense post‐settlement stands. Main conclusions Several lines of evidence suggest that although species composition may have been altered since settlement, the same basic ecosystems occurred on this landscape in pre‐settlement forests, providing reference information for ecological restoration. Red cinders/Bahia ecosystems were rare historically and > 30% of their area has been burned by crown fires since 1950, indicating that priority could be given to restoring this ecosystem's remaining mapping units. Ecosystem classifications may be useful as data layers in gap analyses to identify restoration and conservation priorities. Ecosystem turnover occurs at broad extents on this landscape, and restoration must accordingly operate across large areas to encompass ecosystem diversity. By incorporating factors driving ecosystem composition, this ecosystem classification represents a framework for estimating spatial variation in ecological properties, such as species diversity, relevant to ecological restoration.     

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     

Tree growth response to drought and temperature in a mountain landscape in northern Arizona, USA

Aim To understand how tree growth response to regional drought and temperature varies between tree species, elevations and forest types in a mountain landscape. Location Twenty‐one sites on an elevation gradient of 1500 m on the San Francisco Peaks, northern Arizona, USA. Methods Tree‐ring data for the years 1950–2000 for eight tree species (Abies lasiocarpa var. arizonica (Merriam) Lemm., Picea engelmannii Parry ex Engelm., Pinus aristata Engelm., Pinus edulis Engelm., Pinus flexilis James, Pinus ponderosa Dougl. ex Laws., Pseudotsuga menziesii var. glauca (Beissn.) Franco and Quercus gambelii Nutt.) were used to compare sensitivity of radial growth to regional drought and temperature among co‐occurring species at the same site, and between sites that differed in elevation and species composition. Results For Picea engelmannii, Pinus flexilis, Pinus ponderosa and Pseudotsuga menziesii, trees in drier, low‐elevation stands generally had greater sensitivity of radial growth to regional drought than trees of the same species in wetter, high‐elevation stands. Species low in their elevational range had greater drought sensitivity than co‐occurring species high in their elevational range at the pinyon‐juniper/ponderosa pine forest ecotone, ponderosa pine/mixed conifer forest ecotone and high‐elevation invaded meadows, but not at the mixed conifer/subalpine forest ecotone. Sensitivity of radial growth to regional drought was greater at drier, low‐elevation compared with wetter, high‐elevation forests. Yearly growth was positively correlated with measures of regional water availability at all sites, except high‐elevation invaded meadows where growth was weakly correlated with all climatic factors. Yearly growth in high‐elevation forests up to 3300 m a.s.l. was more strongly correlated with water availability than temperature. Main conclusions Severe regional drought reduced growth of all dominant tree species over a gradient of precipitation and temperature represented by a 1500‐m change in elevation, but response to drought varied between species and stands. Growth was reduced the most in drier, low‐elevation forests and in species growing low in their elevational range in ecotones, and the least for trees that had recently invaded high‐elevation meadows. Constraints on tree growth from drought and high temperature are important for high‐elevation subalpine forests located near the southern‐most range of the dominant species.     

Are differences in productivity between native and exotic trees in N.W. Patagonia related to differences in hydraulic conductance?

Afforestation with the exotic Pinus ponderosa is currently taking place within the natural distributional area of Austrocedrus chilensis, a native conifer of N.W. Patagonia. Annual productivity of the exotic species is double the productivity of the native one. In order to test the hypothesis that these differences in productivity are, at least in part, due to differences in hydraulic characteristics of both species, we measured or estimated several ecophysiological variables in A. chilensis and P. ponderosa trees growing in the same place. Water use (WU) and diameter growth were lower in A. chilensis than in P. ponderosa. Although predawn water potential was relatively constant during the whole growing season, A. chilensis trees showed lower values of this variable than P. ponderosa in a very dry period, suggesting different water sources. Under field conditions, canopy-stomatal (gs) and whole hydraulic conductances, specific hydraulic conductivity and photosynthetic rate (A) were lower in A. chilensis than P. ponderosa. In contrast, instantaneous WU efficiency was higher in A. chilensis than in P. ponderosa. However, gs and A in A. chilensis significantly increased in cut branches of this species suggesting hydraulic limitations on photosynthesis. We hypothesize that hydraulic characteristics of P. ponderosa permit high stomatal conductance for more hours a day than A. chilensis trees, without reaching threshold values of water potential. This can explain, at least in part, differences in C fixation and thus, in productivity between species. In addition, our results suggested a secondary limitation to C fixation in A. chilensis at the photosynthetic apparatus.     

Responses of exotic plant species to fires in Pinus ponderosa forests in northern Arizona

Abstract. Changes in disturbance due to fire regime in southwestern Pinus ponderosa forests over the last century have led to dense forests that are threatened by widespread fire. It has been shown in other studies that a pulse of native, early‐seral opportunistic species typically follow such disturbance events. With the growing importance of exotic plants in local flora, however, these exotics often fill this opportunistic role in recovery. We report the effects of fire severity on exotic plant species following three widespread fires of 1996 in northern Arizona P. ponderosa forests. Species richness and abundance of all vascular plant species, including exotics, were higher in burned than nearby unburned areas. Exotic species were far more important, in terms of cover, where fire severity was highest. Species present after wildfires include those of the pre‐disturbed forest and new species that could not be predicted from above‐ground flora of nearby unburned forests.     

A 40,000-year woodrat-midden record of vegetational and biogeographical dynamics in north-eastern Utah, USA

Aim A conspicuous climatic and biogeographical transition occurs at 40–45° N in western North America. This pivot point marks a north–south opposition of wet and dry conditions at interannual and decadal time‐scales, as well as the northern and southern limits of many dominant western plant species. Palaeoecologists have yet to focus on past climatic and biotic shifts along this transition, in part because it requires comparisons across dissimilar records [i.e. pollen from lacustrine sediments to the north and plant macrofossils from woodrat (Neotoma) middens to the south]. To overcome these limitations, we are extending the woodrat‐midden record northward into the lowlands of the central Rocky Mountains. Location Woodrat middens were collected from crevices and rock shelters on south‐facing slopes of Dutch John Mountain (2000–2200 m, 40°57′ N, 109°25′ W), situated on the eastern flanks of the Uinta Mountains in north‐eastern Utah. The site is near the regional limits for Pinus ponderosa, P. edulis, P. contorta, Cercocarpus ledifolius var. intricatus, Abies concolor, Ephedra viridis and other important western species. Methods We analysed pollen and plant macrofossils from the 40,000‐year midden sequence. The middens represent brief, depositional episodes (mostly years to decades). Four middens represent the early to full‐glacial period (40,000–18,000 cal‐yr bp ), eight middens are from the late‐glacial/early Holocene transition (13,500–9000 cal yr bp ), and 33 middens span the mid‐to‐late Holocene (last 7500 years). Temporal density of our Holocene middens (one every c. 210 years) is comparable with typical Holocene pollen sequences from lake sediments. Results Early to full‐glacial assemblages are characterized by low diversity and occurrence of montane conifers (Picea pungens, Pseudotsuga menziesii, P. flexilis, Juniperus communis) absent from the site today. Diversity increases in the late‐glacial samples with the addition of J. scopulorum, J. horizontalis, C. montanus, C. ledifolius var. intricatus and mesic understory species. The coniferous trees and J. communis declined and J. osteosperma appeared during the late‐glacial/Holocene transition. Juniperus osteosperma populations have occupied the site throughout the Holocene. Pinus ponderosa was established by 7500 cal‐yr bp , and has occurred at least locally ever since. Montane conifers and J. horizontalis persisted until c. 5500 cal‐yr bp . The signature events of the late Holocene were the invasions of P. edulis and Ephedra viridis and establishment of pinyon–juniper woodland in the last 800 years. Main conclusions The Dutch John Mountain midden record adds to an emerging picture in which mid‐elevation conifers (P. flexilis, Pseudotsuga menziesii, Picea pungens, J. scopulorum, J. communis) dominated vegetation over a wide area of the Colorado Plateau and adjacent Rocky Mountains. Rather than being fragmented, as often assumed in phylogeographical studies, these species had broader and more‐connected distributions than they do in the region today. Paradoxically, subalpine conifers (Picea engelmannii, A. lasiocarpa) occurred at higher elevations to the south, possibly representing declining precipitation from south to north owing to southward displacement of the polar jet stream. The Dutch John Mountain record displays a series of extinction and invasion events. Most of the extinctions were local in scale; nearly all constituents of fossil midden assemblages occur within a few kilometres of Dutch John Mountain, and some occur at least locally on its slopes. The sole exception is J. horizontalis, which is regionally extinct. In contrast to extinctions, Holocene invasions were regional in scale; J. osteosperma, P. ponderosa, P. edulis and Ephedra viridis immigrated from glacial‐age source populations far to the south.     

Primers designed to amplify a mitochondrial nad1 intron in ponderosa pine, Pinus ponderosa, limber pine, P. flexilis, and Scots pine, P. sylvestris

The b/c intron of the mitochondrial nad1 gene, was sequenced to characterize the indel region of ponderosa pine, Pinus ponderosa. The sequence in ponderosa pine was aligned with the sequence in Scots pine, Pinus sylvestris, to design seven primers that are useful for sequencing and for revealing size variation in amplified fragments in ponderosa pine, Scots pine, and limber pine, Pinus flexilis. These primers reveal variability in all three species, and the pattern of variability within ponderosa pine is described by a preliminary survey. The indel region of ponderosa pine contains three distinct elements with lengths of 31, 32, and 34 bp. Received: 1 March 2000 / Accepted: 14 April 2000<@head-com-p1a.lf>Communicated by P.M.A. Tigerstedt     

Determining the factors affecting the distribution of Muscari latifolium,an endemic plant of Turkey,and a mapping species distribution model

Species distribution modeling was used to determine factors among the large predictor candidate data set that affect the distribution of Muscari latifolium , an endemic bulbous plant species of Turkey, to quantify the relative importance of each factor and make a potential spatial distribution map of M. latifolium . Models were built using the Boosted Regression Trees method based on 35 presence and 70 absence records obtained through field sampling in the Gönen Dam watershed area of the Kazda?? Mountains in West Anatolia. Large candidate variables of monthly and seasonal climate, fine‐scale land surface, and geologic and biotic variables were simplified using a BRT simplifying procedure. Analyses performed on these resources, direct and indirect variables showed that there were 14 main factors that influence the species’ distribution. Five of the 14 most important variables influencing the distribution of the species are bedrock type, Quercus cerris density, precipitation during the wettest month, Pinus nigra density, and northness. These variables account for approximately 60% of the relative importance for determining the distribution of the species. Prediction performance was assessed by 10 random subsample data sets and gave a maximum the area under a receiver operating characteristic curve (AUC) value of 0.93 and an average AUC value of 0.8. This study provides a significant contribution to the knowledge of the habitat requirements and ecological characteristics of this species. The distribution of this species is explained by a combination of biotic and abiotic factors. Hence, using biotic interaction and fine‐scale land surface variables in species distribution models improved the accuracy and precision of the model. The knowledge of the relationships between distribution patterns and environmental factors and biotic interaction of M. latifolium can help develop a management and conservation strategy for this species.     

Associations Among Breeding Birds and Gambel Oak in Southwestern Ponderosa Pine Forests

Abstract: Ponderosa pine (Pinus ponderosa) forests with Gambel oak (Quercus gambelii) are associated with higher bird abundance and diversity than are ponderosa pine forests lacking Gambel oak. Little is known, however, about specific structural characteristics of Gambel oak trees, clumps, and stands that may be important to birds in ponderosa pine-Gambel oak (hereafter pine-oak) forests. We examined associations among breeding birds and structural characteristics of Gambel oak at a scale similar in size to individual bird territories in pine-oak forests in northern Arizona and western New Mexico, USA. Avian species richness and occurrence of some bird species were associated with specific growth forms of Gambel oak. Estimated probability of Virginia's warblers (Vermivora virginiae), black-headed grosbeaks (Pheucticus melanocephalus), and red-faced warblers (Cardellina rubrifrons) occurring at points increased with increasing density of pole-sized Gambel oak 7–15 cm in diameter at breast height. We also found evidence that large Gambel oak trees (≥23 cm dbh) were associated with increased occurrence of yellow-rumped warblers (Dendroica coronata) at points. Some avian associations with oak were influenced by characteristics of ponderosa pines. For example, bird species richness was positively associated with the abundance of large Gambel oak when density of large pine trees ≥23 cm in diameter at breast height was low. Because large oak trees are rare and their numbers are thought to be declining, efforts should be made to retain and promote growth of additional oaks in this size class. Forest management practices that maintain forest openings, such as prescribed burning, could promote growth of pole-sized Gambel oak, which appears important to some bird species in pine-oak forests.     

Ectomycorrhizal communities of ponderosa pine and lodgepole pine in the south-central Oregon pumice zone

Forest ecosystems of the Pacific Northwest of the USA are changing as a result of climate change. Specifically, rise of global temperatures, decline of winter precipitation, earlier loss of snowpack, and increased summer drought are altering the range of Pinus contorta. Simultaneously, flux in environmental conditions within the historic P. contorta range may facilitate the encroachment of P. ponderosa into P. contorta territory. Furthermore, successful pine species migration may be constrained by the distribution or co-migration of ectomycorrhizal fungi (EMF). Knowledge of the linkages among soil fungal diversity, community structure, and environmental factors is critical to understanding the organization and stability of pine ecosystems. The objectives of this study were to establish a foundational knowledge of the EMF communities of P. ponderosa and P. contorta in the Deschutes National Forest, OR, USA, and to examine soil characteristics associated with community composition. We examined EMF root tips of P. ponderosa and P. contorta in soil cores and conducted soil chemistry analysis for P. ponderosa cores. Results indicate that Cenococcum geophilum, Rhizopogon salebrosus, and Inocybe flocculosa were dominant in both P. contorta and P. ponderosa soil cores. Rhizopogon spp. were ubiquitous in P. ponderosa cores. There was no significant difference in the species composition of EMF communities of P. ponderosa and P. contorta. Ordination analysis of P. ponderosa soils suggested that soil pH, plant-available phosphorus (Bray), total phosphorus (P), carbon (C), mineralizable nitrogen (N), ammonium (NH₄), and nitrate (NO₃) are driving EMF community composition in P. ponderosa stands. We found a significant linear relationship between EMF species richness and mineralizable N. In conclusion, P. ponderosa and P. contorta, within the Deschutes National Forest, share the same dominant EMF species, which implies that P. ponderosa may be able to successfully establish within the historic P. contorta range and dominant EMF assemblages may be conserved.     

GENETIC VARIATION IN THE PONDEROSAE OF THE SOUTHWEST

Ninety-five seedling populations of southwestern ponderosa pine (Pinus ponderosa var. scopulorum) along with single populations of Pinus engelmannii and Pinus arizonica were compared in four environmentally disparate common gardens. Differentiation among ponderosa pine populations was detected for a diverse assortment of variables that included patterns of shoot elongation, measures of growth potential, winter and spring freezing damage, and leaf characteristics. Multiple regression models accounted for as much as 85% of the variance among populations and described complex clines that were dominated by elevational and latitudinal effects. Although P. ponderosa, P. arizonica, and P. engelmannii were readily differentiated, the performance of progenies from one population suggested introgression primarily involving P. ponderosa and P. arizonica but also implicating P. engelmannii.     

The 5S RNA genes inPinus radiata and the spacer region as a probe for relationships betweenPinus species

The 5S RNA genes inPinus radiata occur in two size classes of about 850 and 525 bp in length. Representatives from both the long and short size classes were cloned and sequenced. The primary difference in the two size classes was shown to be a 330 bp insertion in the spacer region of the long units. Within an individual breeding clone ofP. radiata there was some sequence heterogeneity between representatives of the short class. The 5S RNA genes in thirty pine species were characterised using either a clone of the short size class or a subclone of the 330 bp insertion characterizing the long size class. Eleven species of subg.Strobus were examined and all lacked the long type of unit of radiata pine. The New World species of subg.Pinus all had both short and long units whereas the Old World species had long units. The implications of these results for the evolution of the 5S DNA sequences and the phylogenetic relationships withinPinus are discussed.     

Radial growth responses to gap creation in large,old Sequoiadendron giganteum

Questions: Do large, old Sequoiadondron giganteum trees respond to the creation of adjacent canopy gaps? Do other co‐occurring tree species and younger S. giganteum adjacent to gaps also respond? What are the likely factors affecting growth responses? Location: Mixed‐conifer forests of the southern Sierra Nevada, California, USA. Methods: We measured the growth response of large, old S. giganteum trees (mean DBH=164 cm; ages estimated >1000 yr) to gap creation by coring trees and comparing growth after gap creation to growth before gap creation. We also measured young Abies concolor, Pinus lambertiana, and young S. giganteum. Gap‐adjacent trees were compared with non‐adjacent reference trees. Tree rings were analysed for carbon isotope discrimination and for longer‐term growth trend correlations with climate. Results: Following gap creation gap‐adjacent old S. giganteum grew more than reference trees. Abies concolor trees also exhibited a growth response to gap creation. No response was detected for young S. giganteum or P. lambertiana, although detection power was lower for these groups. There was no difference in carbon isotope discrimination response to gap creation between gap‐adjacent and reference trees for old S. giganteum and radial growth was positively correlated with winter precipitation, but not growing season temperature. Conclusion: It is unclear what caused the growth release in old S. giganteum trees, although liberation of below‐ground resources following removal of competing vegetation appears to be a significant contributor. Sequoiadondron giganteum, the third‐longest lived and the largest of all species, remains sensitive to local environmental changes even after canopy emergence. Management activities that reduce vegetation surrounding individual specimen trees can be expected to result in increased vigor of even these very old and large trees.     

Photosynthetic performance of invasive Pinus ponderosa and Juniperus virginiana seedlings under gradual soil water depletion

Changes in climate, land management and fire regime have contributed to woody species expansion into grasslands and savannas worldwide. In the USA, Pinus ponderosa P.&C. Lawson and Juniperus virginiana L. are expanding into semiarid grasslands of Nebraska and other regions of the Great Plains. We examined P. ponderosa and J. virginiana seedling response to soil water content, one of the most important limiting factors in semiarid grasslands, to provide insight into their success in the region. Photosynthesis, stomatal conductance, maximum photochemical efficiency of PSII, maximum carboxylation velocity, maximum rate of electron transport, stomatal limitation to photosynthesis, water potential, root‐to‐shoot ratio, and needle nitrogen content were followed under gradual soil water depletion for 40 days. J. virginiana maintained lower L_s, higher A, g_s, and initial F_v/F_m, and displayed a more gradual decline in V_cmax and J_max with increasing water deficit compared to P. ponderosa. J. virginiana also invested more in roots relative to shoots compared to P. ponderosa. F_v/F_m showed high PSII resistance to dehydration in both species. Photoinhibition was observed at ～30% of field capacity. Soil water content was a better predictor of A and g_s than Ψ, indicating that there are other growth factors controlling physiological processes under increased water stress. The two species followed different strategies to succeed in semiarid grasslands. P. ponderosa seedlings behaved like a drought‐avoidant species with strong stomatal control, while J. virginiana was more of a drought‐tolerant species, maintaining physiological activity at lower soil water content. Differences between the studied species and the ecological implications are discussed.     

Factors in Pinus ponderosa and Calocedrus decurrens mortality in Yosemite Valley,USA

Management practices over the past 100 years, especially fire exclusion and suppression, in Yosemite Valley, Yosemite National Park, CA resulted in forest stands with basal areas in excess of 110 m² ha^-1. Accelerated attack by several species of bark beetles (Scolytidae) followed a severe drought in 1976–77. Pinus ponderosa, already under attack by root rot (Heterobasidion annosus) was affected severely, and mortality exceeded 70%. Calocedrus decurrens did not suffer comparable mortality and has replaced P. ponderosa as the dominant species. Size class distributions as well as comparisons of relative density and relative basal area from 1961, 1976 and 1983–85 also demonstrate a major change in forest composition. This change is most pronounced on alluvial landforms.     

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.