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.     

Drought responses of conifers in ecotone forests of northern Arizona: tree ring growth and leaf δ<Superscript>13</Superscript>C

We sought to understand differences in tree response to meteorological drought among species and soil types at two ecotone forests in northern Arizona, the pinyon-juniper woodland/ponderosa pine ecotone, and the higher elevation, wetter, ponderosa pine/mixed conifer ecotone. We used two approaches that provide different information about drought response: the ratio of standardized radial growth in wet years to dry years (W:D) for the period between years 1950 and 2000 as a measure of growth response to drought, and ¹³C in leaves formed in non-drought (2001) and drought (2002) years as a measure of change in water use efficiency (WUE) in response to drought. W:D and leaf ¹³C response to drought for Pinus edulis and P. ponderosa did not differ for trees growing on coarse-texture soils derived from cinders compared with finer textured soils derived from flow basalts or sedimentary rocks. P. ponderosa growing near its low elevation range limit at the pinyon-juniper woodland/ponderosa pine ecotone had a greater growth response to drought (higher W:D) and a larger increase in WUE in response to drought than co-occurring P. edulis growing near its high elevation range limit. P. flexilis and Pseudotsuga menziesii growing near their low elevation range limit at the ponderosa pine/mixed conifer ecotone had a larger growth response to drought than co-occurring P. ponderosa growing near its high elevation range limit. Increases in WUE in response to drought were similar for all species at the ponderosa pine/mixed conifer ecotone. Low elevation populations of P. ponderosa had greater growth response to drought than high-elevation populations, whereas populations had a similar increase in WUE in response to drought. Our findings of different responses to drought among co-occurring tree species and between low- and high-elevation populations are interpreted in the context of drought impacts on montane coniferous forests of the southwestern USA.     

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.     

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     

Habitat Conditions Associated With Lynx Hunting Behavior During Winter in Northern Washington

Abstract Effectively managing habitat for threatened populations of Canada lynx (Lynx canadensis) requires knowledge of habitat conditions that provide for the ecological needs of lynx. We snow-tracked lynx to identify habitat conditions associated with hunting behavior and predation during winters of 2002–2003 and 2003–2004 in the northern Cascade Range in Washington state, USA. We recorded number and success of predation attempts, prey species killed, and trail sinuosity on 149 km of lynx trails. Lynx killed snowshoe hares (Lepus americanus), red squirrels (Tamiasciurus hudsonicus), and cricetids more than expected in Englemann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests, where snowshoe hare densities were highest. Lynx killed prey less than expected in Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) forests and forest openings. We used the sinuosity of lynx trails as an index of quality of habitat hunted. Lynx trails that included predation attempts were more sinuous than trail segments without predation attempts. Lynx trails had greater sinuosity in forest stands with high hare densities dominated by Engelmann spruce and subalpine fir than in stands with low hare densities dominated by Douglas-fir and ponderosa pine or in forest openings. We encourage forest managers to maintain or create sufficient understory cover to support high densities of snowshoe hares as foraging habitat for lynx.     

GENETIC STRUCTURE OF GEOGRAPHICALLY MARGINAL POPULATIONS OF PONDEROSA PINE

Eleven populations of ponderosa pine (Pinus ponderosa) located on an east-west transect of the Palmer Divide of Central Colorado were studied. Within each population transects of approximately 20 × 250 m were established. In seven populations the transects represented contrasting ecological conditions such as north and south facing slopes. Within each transect 200–250 trees greater than 10 cm in height were tagged, their locations mapped, their ages determined and their diameters and heights measured. Electrophoretic analyses of mature needle tissue indicated that while most of the allozyme variation resided within transects, significant genetic heterogeneity could be found between populations and transects. Among populations consistent patterns of genetic variation were observed at only two loci. Five of the 13 loci had consistent but very small differences between transects within populations. Considerable genetic heterogeneity occurred among 20 m sections of the transects and among age classes within transects. The genetic structure of these populations can be explained by mosaic patterns of seedling recruitment characteristic of these ponderosa pine stands. Only a few maternal trees contribute to each patch of seedlings, and this localized founder effect produces the genetic structure observed in these populations.     

EXPERIMENTAL STUDIES OF PONDEROSA PINE. III. DIFFERENCES IN PHOTOSYNTHESIS,STOMATAL CONDUCTANCE,AND WATER-USE EFFICIENCY BETWEEN TWO GENETIC LINES

As part of an intensive study of heritable differences among the progeny of Pinus ponderosa parents from two contrasting habitats (coastal vs. interior, continental), we examined the potential for differences in photosynthesis rate, stomatal conductance, and photosynthetic water-use efficiency. Plants from a cross between two coastal parents (ponderosa × ponderosa) exhibited lower photosynthetic water-use efficiencies, relative to plants from a coastal × interior cross (ponderosa × scopulorum). The lower water-use efficiencies in the ponderosa × ponderosa plants were evident as a lower ratio of external to intercellular CO₂ concentrations and higher stomatal conductances at any given rate of photosynthesis. The ponderosa × scopulorum plants exhibited lower stomatal conductances over a range of leaf-to-air water vapor concentration differences, which was partially explained by lower stomatal densities. The ponderosa × scopulorum plants also exhibited lower maximum photosynthesis rates and lower needle nitrogen concentrations. Taken together, the results suggest that in adapting to drier habitats, P. ponderosa has acquired improved water-use efficiencies and lower transpiration rates, but at the expense of reduced maximum photosynthesis rates.     

Classification tree and minimum-volume ellipsoid analyses of the distribution of ponderosa pine in the western USA

Aim Ponderosa pine (Pinus ponderosa Douglas ex Lawson & C. Lawson) is an economically and ecologically important conifer that has a wide geographic range in the western USA, but is mostly absent from the geographic centre of its distribution – the Great Basin and adjoining mountain ranges. Much of its modern range was achieved by migration of geographically distinct Sierra Nevada (P. ponderosa var. ponderosa) and Rocky Mountain (P. ponderosa var. scopulorum) varieties in the last 10,000 years. Previous research has confirmed genetic differences between the two varieties, and measurable genetic exchange occurs where their ranges now overlap in western Montana. A variety of approaches in bioclimatic modelling is required to explore the ecological differences between these varieties and their implications for historical biogeography and impending changes in western landscapes. Location Western USA. Methods We used a classification tree analysis and a minimum‐volume ellipsoid as models to explain the broad patterns of distribution of ponderosa pine in modern environments using climatic and edaphic variables. Most biogeographical modelling assumes that the target group represents a single, ecologically uniform taxonomic population. Classification tree analysis does not require this assumption because it allows the creation of pathways that predict multiple positive and negative outcomes. Thus, classification tree analysis can be used to test the ecological uniformity of the species. In addition, a multidimensional ellipsoid was constructed to describe the niche of each variety of ponderosa pine, and distances from the niche were calculated and mapped on a 4‐km grid for each ecological variable. Results The resulting classification tree identified three dominant pathways predicting ponderosa pine presence. Two of these three pathways correspond roughly to the distribution of var. ponderosa, and the third pathway generally corresponds to the distribution of var. scopulorum. The classification tree and minimum‐volume ellipsoid model show that both varieties have very similar temperature limitations, although var. ponderosa is more limited by the temperature extremes of the continental interior. The precipitation limitations of the two varieties are seasonally different, with var. ponderosa requiring significant winter moisture and var. scopulorum requiring significant summer moisture. Great Basin mountain ranges are too cold at higher elevations to support either variety of ponderosa pine, and at lower elevations are too dry in summer for var. scopulorum and too dry in winter for var. ponderosa. Main conclusions The classification tree analysis indicates that var. ponderosa is ecologically as well as genetically distinct from var. scopulorum. Ecological differences may maintain genetic separation in spite of a limited zone of introgression between the two varieties in western Montana. Two hypotheses about past and future movements of ponderosa pine emerge from our analyses. The first hypothesis is that, during the last glacial period, colder and/or drier summers truncated most of the range of var. scopulorum in the central Rockies, but had less dramatic effects on the more maritime and winter‐wet distribution of var. ponderosa. The second hypothesis is that, all other factors held constant, increasing summer temperatures in the future should produce changes in the distribution of var. scopulorum that are likely to involve range expansions in the central Rockies with the warming of mountain ranges currently too cold but sufficiently wet in summer for var. scopulorum. Finally, our results underscore the growing need to focus on genotypes in biogeographical modelling and ecological forecasting.     

Influence of induced water stress in ponderosa pine on pine sawflies

Summary Three levels of water stress were induced on pole-size ponderosa pine (Pinus ponderosa) to determine the influence of plant moisture stress on oviposition, survival, and growth of two species of pine sawfly (Neodiprion fulviceps and N. autumnalis). It was found that water stress affected oviposition and subsequent egg survival but not larval development or survival. Stress had a negative effect on early season oviposition (N. fulviceps) and a positive effect on late season oviposition (N. autumnalis). Egg hatch was different between species and years and among treatment levels. Larval development, feeding, and survival were not affected by water stress. Overall, the effect of stress was not sufficient to explain population outbreaks of sawflies. Several hypotheses are presented as possible explanations for the observed experimental results.     

Observations on the genetic structure and mating system of ponderosa pine in the Colorado front range

Summary Variation of peroxidase enzymes is analyzed both in mature needle tissue and in open-pollinated seedling families of ponderosa pine, Pinus ponderosa, and is identified as being controlled by a single Mendelian locus. Variation at this locus, analyzed in 1, 386 individuals, is used in the analysis of population differentiation and the mating system. Significant variation of gene frequencies is detected over distances of several hundred meters, and is found to be associated with slopes of different aspects. Ponderosa pine is wind-pollinated, and an analysis of the mating system indicates that the level of outcrossing is greater than 90 %. Selection specific for different environments is evidently strong enough to overcome the homogenizing force of migration and produce population fissuring in ponderosa pine.     

Roosts of Allen's Lappet-Browed Bat in Northern Arizona

ABSTRACT In Arizona, USA, Allen's lappet-browed bat (Idionycteris phyllotis) forms maternity colonies in ponderosa pine (Pinus ponderosa) snags. There is little information on the roosting habitat of males. We used radiotelemetry to locate 16 maternity, 3 postlactating, and 2 bachelor roosts and combined data with unpublished data for maternity roosts (n = 11) located in 1993–1995. Most (96%) maternity roosts were in large-diameter (x̄ ± SE: 64 ± 2.7 cm) ponderosa pine snags under sloughing bark. Models that best predicted the probability of a snag's use as a maternity roost indicated bats selected taller snags closer to forest roads than comparison snags. Maternity roosts averaged 11 bats per roost (SE = 2, n = 15; from exit counts) and were an average distance of 1.6 km from capture sites (SE = 0.3, n = 17). Bachelor roosts were in vertical sandstone cliff faces in pinyon-juniper (Pinus edulis-Juniperus spp.) woodlands approximately 12 km from capture sites; these and other capture records in Arizona indicated sexual segregation may have occurred during the maternity season. Of 11 maternity snag roosts located in 1993–1995, only one continued to function as a roost. Resource managers should maintain patches of large-diameter ponderosa pine snags with peeling bark to provide maternity roosting habitat for Allen's lappet-browed bat.     

Persistence and expansion of ponderosa pine woodlands in the west‐central Great Plains during the past two centuries

Aim Woody plant expansion and infilling in grasslands and savannas are occurring across a broad range of ecosystems around the globe and are commonly attributed to fire suppression, livestock grazing, nutrient enrichment and/or climate variability. In the western Great Plains, ponderosa pine (Pinus ponderosa) woodlands are expanding across broad geographical and environmental gradients. The objective of this study was to reconstruct the establishment of ponderosa pine in woodlands in the west‐central Great Plains and to identify whether it was mediated by climate variability. Location Our study took place in a 400‐km wide region from the base of the Front Range Mountains (c. 105° W) to the central Great Plains (c. 100° W) and from Nebraska (43° N) to northern New Mexico (36° N), USA. Methods Dates for establishment of ponderosa pine were reconstructed with tree rings in 11 woodland sites distributed across the longitudinal and latitudinal gradients of the study area. Temporal trends in decadal pine establishment were compared with summer Palmer Drought Severity Index (PDSI). Annual trends in pine establishment from 1985 to 2005 were compared with seasonal PDSI, temperature and moisture availability. Results Establishment of ponderosa pine occurred in the study area in all but one decade (1770s) between the 1750s and the early 2000s, with over 35% of establishment in the region occurring after 1980. Pine establishment was highly variable among sites. Across the region, decadal pine establishment was persistently low from 1940 to 1960, when PDSI was below average. Annual pine establishment from 1985 to 2005 was positively correlated with summer PDSI and inversely correlated with minimum spring temperatures. Main conclusions Most ponderosa pine woodlands pre‐date widespread Euro‐American settlement of the region around c. ad 1860 and currently have stable tree populations. High variability in the timing of establishment of pine among sites highlights the multiplicity of factors that can drive woodland dynamics, including land use, fire history, CO₂ enrichment, tree population dynamics and climate. Since the 1840s, the influence of climate was most notable across the study area during the mid‐20th century, when the establishment of pine was suppressed by two significant droughts. The past sensitivity of establishment of ponderosa pine to drought suggests that woodland expansion will be negatively affected by predicted increases in temperature and drought in the Great Plains.     

Disturbance and climatic influences on age structure of ponderosa pine at the pine/grassland ecotone, Colorado Front Range

Abstract. In the Colorado Front Range, disturbances and climatic variation influence stand structure of ponderosa pine (Pinus ponderosa) along the lower timberline ecotone. Over the past 100 years there has been a shift to a greater density and extent of ponderosa pine at the forest-grassland boundaries. Ponderosa pine regeneration at lower timberline appears to be influenced by fires in the 1860s and decreased grazing pressure in the 1970s–1980s. Climatic variation may also have influenced age structure, even though analyses of age structure at a 10-year class scale prevented the detection of climatic influences occurring at a finer scale. These changes in disturbance regimes, possibly together with moister springs/early summers, created favourable conditions for the increase in density and extent of ponderosa pine at lower timberline ecotone.     

Climate-related trends in sapwood biophysical properties in two conifers: avoidance of hydraulic dysfunction through coordinated adjustments in xylem efficiency, safety and capacitance

In the Pacific north‐west, the Cascade Mountain Range blocks much of the precipitation and maritime influence of the Pacific Ocean, resulting in distinct climates east and west of the mountains. The current study aimed to investigate relationships between water storage and transport properties in populations of Douglas‐fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) adapted to both climates. Sapwood thickness, capacitance, vulnerability to embolism, and axial and radial conductivity were measured on samples collected from trunks of mature trees. The sapwood of ponderosa pine was three to four times thicker than Douglas‐fir. Radial conductivity was higher in west‐side populations of both species, but axial conductivity was higher in the east‐side populations and in Douglas‐fir. Eastern populations of both species had sapwood that was more vulnerable to embolism than west‐side populations. Sapwood capacitance was similar between species, but was about twice as great in east‐side populations (580 kg m^?3 MPa^?1) as in west‐side populations (274 kg m^?3 MPa^?1). Capacitance was positively correlated with both mean embolism pressure and axial conductivity across species and populations, suggesting that coordinated adjustments in xylem efficiency, safety and water storage capacity may serve to avoid embolism along a gradient of increasing aridity.     

GENETIC PATTERNS ASSOCIATED WITH GROWTH VARIABILITY IN PONDEROSA PINE

Variability in annual growth increment in Pinus ponderosa was measured and heterozygosity level estimated in order to examine the relationship between genetic heterozygosity and phenotypic homeostasis in natural populations of a long-lived perennial plant. Pollen was collected from 190 ponderosa pine trees in the upper montane region of the Front Range in Colorado and four isozyme systems were resolved electrophoretically from these samples. Each tree was cored, aged, and each annual ring width measured. Statistics estimating within-tree variation in growth increment were then related to heterozygosity levels. The results indicated a significant positive association between level of heterozygosity and growth variability. Two speculations are offered to account for this relationship.     

ALLELOPATHIC INHIBITION OF NITRIFICATION AND NITRIFYING BACTERIA IN A PONDEROSA PINE (PINUS PONDEROSA DOUGL.) COMMUNITY

The dynamics of ponderosa pine (Pinus ponderosa Dougl.) stands in western North Dakota were studied to determine the influence of plant-produced chemicals on nitrification rates and competitive interactions within the stands. Ponderosa pine accounted for more than 98% of all tree and shrub stratum stems in this climax community. Low levels of nitrate-nitrogen relative to ammonium-nitrogen and low numbers of Nitrosomonas and Nitrobacter in the soils indicated that nitrification rates were low. Inhibition of nitrification is often attributed to low soil pH in coniferous forests, but the slightly alkaline soils in this study (pH 7.25–7.75) suggested that another factor caused the low nitrification. Evidence obtained suggested that the reduction in nitrate synthesis was due to the production and subsequent transfer to the soil of secondary plant chemicals that were toxic to Nitrosomonas. Chemical inhibitors of nitrification, including caffeic acid, chlorogenic acid, quercitin, and condensed tannins, were found in extracts from ponderosa pine needles, bark, and A horizon soils. These extracts proved to be toxic to soil suspensions of Nitrosomonas causing reductions of from 68–93% of the control. These findings indicate that climax ponderosa pine communities minimize the conversion of ammonia-nitrogen to nitrate-nitrogen by chemically inhibiting nitrification.     

Water and Nutrient Outflow Following the Ecological Restoration of a Ponderosa Pine-Bunchgrass Ecosystem

In the late 1800s, fire suppression, livestock grazing, and a wet and warm climate led to an irruption of pine regeneration in Pinus ponderosa Laws. (ponderosa pine) forests of the southwestern United States. Pines invaded bunchgrass openings, causing stand structure changes that increased the number of stand-replacing fires. Ecological restoration, via thinning and prescribed burning, is being used to decrease the risk of stand-replacing fires and ameliorate other effects of pine invasion. The effects of aboveground restoration on belowground processes are poorly understood. We used a hydrologic model and soil water nutrient concentrations, measured monthly below the rooting zone, to estimate restoration effects on nutrient losses by leaching from a mature ponderosa pine forest near Flagstaff, Arizona. Replicated restoration treatments included thinning to pre-1880 stand densities (partial restoration), thinning plus forest floor fuel reduction followed by a prescribed burn (complete restoration), and an untreated control. Water outflow occurred only between January and May and was lowest from the control (47 and 28 mm in 1995 and 1996) and highest from the partial restoration treatment (67 and 59 mm in 1995 and 1996). The concentrations (typically <0.10 mg/ L) and estimated annual losses (<0.02 kg/ha) of NH₄⁺-N, PO₄^{3 ?} -P, and organic P were similar among treatments. Nitrate and organic N concentrations were as high as 0.80 mg N/L; however, these concentrations and estimated annual losses (<0.13 kg N/ha) were similar among treatments. Our results suggest that restoration will not enhance nutrient loss by leaching or alter stream chemistry in ponderosa pine forests.     

EXPERIMENTAL STUDIES OF PONDEROSA PINE. II. QUANTITATIVE GENETICS OF MORPHOLOGICAL TRAITS

We studied fourteen morphological attributes of 511 seedling stage progeny involving crosses between Pinus ponderosa ponderosa parents as well as intervarietal crosses of P. p. ponderosa × P. p. scopulorum. Intravarietal progeny were distinctly differentiated genetically from intervarietal sibships in seven of the fourteen characters examined. The patterns of differentiation observed strongly suggest a syndrome of selective responses to increased water stress in the progeny with a scopulorum parent and for comparatively more rapid, columnar growth in the progeny of the intravariety crosses which involved only P. p. ponderosa parents. These differences are consistent with known ecological distinctions between the habitats of the two varieties. Narrow-sense heritability estimates, obtained from half-sib progeny analysis, indicated that considerable levels of additive genetic variance remain present in several traits. We could detect no relationship between the amount of additive genetic variance present for a particular character and its presumed relevance to fitness.     

Exploring Climate Niches of Ponderosa Pine (Pinus ponderosa Douglas ex Lawson) Haplotypes in the Western United States: Implications for Evolutionary History and Conservation

Ponderosa pine (Pinus ponderosa Douglas ex Lawson) occupies montane environments throughout western North America, where it is both an ecologically and economically important tree species. A recent study using mitochondrial DNA analysis demonstrated substantial genetic variation among ponderosa pine populations in the western U.S., identifying 10 haplotypes with unique evolutionary lineages that generally correspond spatially with distributions of the Pacific (P. p. var. ponderosa) and Rocky Mountain (P. p. var. scopulorum) varieties. To elucidate the role of climate in shaping the phylogeographic history of ponderosa pine, we used nonparametric multiplicative regression to develop predictive climate niche models for two varieties and 10 haplotypes and to hindcast potential distribution of the varieties during the last glacial maximum (LGM), ~22,000 yr BP. Our climate niche models performed well for the varieties, but haplotype models were constrained in some cases by small datasets and unmeasured microclimate influences. The models suggest strong relationships between genetic lineages and climate. Particularly evident was the role of seasonal precipitation balance in most models, with winter- and summer-dominated precipitation regimes strongly associated with P. p. vars. ponderosa and scopulorum, respectively. Indeed, where present-day climate niches overlap between the varieties, introgression of two haplotypes also occurs along a steep clinal divide in western Montana. Reconstructed climate niches for the LGM suggest potentially suitable climate existed for the Pacific variety in the California Floristic province, the Great Basin, and Arizona highlands, while suitable climate for the Rocky Mountain variety may have existed across the southwestern interior highlands. These findings underscore potentially unique phylogeographic origins of modern ponderosa pine evolutionary lineages, including potential adaptations to Pleistocene climates associated with discrete temporary glacial refugia. Our predictive climate niche models may inform strategies for further genetic research (e.g., sampling design) and conservation that promotes haplotype compatibility with projected changes in future climate.     

RELATIONSHIP BETWEEN SELF-FERTILITY,ALLOCATION OF GROWTH,AND INBREEDING DEPRESSION IN THREE CONIFEROUS SPECIES

Mortality and growth of self and outcross families of three wind-pollinated, mixed-mating, long-lived conifers, Douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), and noble fir (Abies procera) were followed from outplanting to age 26 (25 for noble fir) in spaced plantings at a common test site. Response to inbreeding differed greatly among species over time and in all regards. Only Douglas-fir and noble fir will be contrasted here, because ponderosa pine usually was intermediate to the other two in its response to inbreeding. In earlier reports, compared to noble fir Douglas-fir had a higher rate of primary selfing and larger inbreeding depression in seed set. Douglas-fir continued to have higher inbreeding depression in nursery and early field survival. The species differed in time courses of inbreeding depression in height and in allocation of growth due to crowding. Between ages 6 and 12, the relative elongation rate (dm · dm^?1 · yr^?1) of Douglas-fir was significantly greater in the selfs than in the outcrosses. The response was not observed in noble fir. At final measurement, inbreeding depression in diameter relative to inbreeding depression in height was greater in Douglas-fir than in noble fir. At final measurement inbreeding depression in height was inversely related to inbreeding depression in survival. Cumulative inbreeding depressions from time of fertilization to final measurement were 0.98, 0.94, and 0.83 for Douglas-fir, ponderosa pine, and noble fir, respectively, which indicates that selfs will not contribute to the mature, reproductive populations.