Variability in the essential oils of wood and foliage of Pinus aristata and Pinus longaeva

Variations in the monoterpenoid composition of the essential oil of wood, and of the mono- and sesquiterpenoids of foliage collected from more than 200 specimens of pines of subsection Balfourianae (P. aristata, P. longaeva and P. balfouriana) substantiate the previously reported morphological and chemical separation of P. aristata and P. longaeva and suggest a restricted intermediacy in northern Arizona and eastern California. On the basis of wood essential oil the limited population of P. aristata in northern Arizona, while exhibiting to a slight degree morphological intermediacy, is found to be practically identical to the main populations of P. aristata in Colorado and New Mexico. On the basis of foliage oil, however, the same population is clearly intermediate, but closer to P. aristata. The detailed chemical data suggest a distinct chemical status for the Arizona population, with selection for the present-day chemophenotypes being the dominant factor in their evolution. Analysis of monoterpenoids from P. balfouriana × longaeva hybrids indicates that incomplete selective elimination of 3-carene-producing P. longaeva chemophenotypes rather than gene exchange with P. balfouriana is responsible for the presence of abnormal P. aristata-like 3-carene-producing trees in eastern California. The latter trees could have resulted from limited gene exchange between P. aristata and P. longaeva by routes no longer available to them lying to the south of the Grand Canyon.     

Spiral grain in bristlecone pines (Pinus longaeva) exhibits no correlation with environmental factors

Six hundred and eleven Great Basin bristlecone pines (Pinus longaeva) were surveyed in two separate groves in California’s White Mountains. The presence and direction of spiral grain were recorded for each tree as well as elevation above sea level, horizon angles, latitude and longitude, trunk diameter, whether the tree was dead, and whether the trunk was broken. The proportions of left-handed, right-handed and straight trees were similar in every part of both groves, although the groves lie at different elevations. No significant correlation was found between the direction of spiral grain and any environmental factor. The hypothesis that spiral grain is an adaptation to distribute sap evenly between the roots and the crown in Pinus longaeva is not strongly supported, since spiral grain is not correlated with asymmetric environments and most trees exhibit <90° rotation through the main stem. The data also do not support the idea that spiral grain makes the tree more resistant to breaking in strong winds. Right-handed spiral grain is predicted by this hypothesis, but most bristlecone pines are either left-handed or exhibit no spiral grain. Bristlecone pines are often uprooted from thin soils by strong winds, but rarely are the main stems broken by this mechanism. Spiral grain in Pinus longaeva growing in California’s White Mountains does not appear to be under environmental control.     

Variability of the wood monoterpenoids from Pinus aristata

Striking differences in turpentine composition between California-Nevada-Utah populations and Colorado-New Mexico-Arizona populations of the Bristlecone pine (Pinus aristata) were demonstrated by GLC analyses of the wood extracts. These differences are in line with Bailey's taxonomic division of Pinus aristata sensu lato into western Pinus longaeva and eastern Pinus aristata sensu stricto.     

Terpenoid and morphological variability of Pinus quadrifolia and the natural hybridization with Pinus monophylla in the San Jacinto Mountains in California

Monoterpenoids from wood of Pinus quadrifolia and Pinus monophylla from south of the San Jacinto Mountains in southern California were analysed by gas-liquid chromatography (GLC). The number of needles per fascicle, the number of resin canals, and the number of abaxial and adaxial stomatal rows in the needles were determined. Percentages of mycrene, α-pinene, and to a minor extent of camphene, β-pinene, limonene, and β-phellandrene, the above mentioned morphological characteristics and previously obtained data were all used for identification and characterisation of species intermediacy in the San Jacinto area. It was concluded that many P. quadrifolia trees in the mixed Buck Ridge stand and some trees in two pure stands of the same area were hybrids and that sympatric and to a lesser extent allopatric introgression of P. monophylla into P. quadrifolia takes place in the San Jacinto region. Introgression of P. quadrifolia into P. monophylla could not be demonstrated.     

On the question of natural hybridization between Pinus discolor and Pinus cembrides

Sixty Pinus cembroides and P. discolor trees from two mixed stands and eight pure stands were analysed for the composition of their wood monoterpenes, number of abaxial and adaxial stomatal rows and the number of needles per fascicle. While by far the majority of the characters analysed showed highly significant differences between the two species, only the sum of thujene, sabinene, γ-terpenene, terpinolene and p-cymene percentages, the sum of myrcene and cis-ocimene percentages, the number of abaxial stomatal rows, and the number of needles per fascicle could be used to evaluate the potential intermediacy of trees from the common sites. The results indicated the absence of any hybridization in the Sierra de la Catarina Mountains of Chihuathua (Mexico) and either a possible minute introgression of P. cembroides into P. discolor in the San Miguelito Mountains of San Luis Potosi (Mexico) or a slightly different genetic composition of the trees from this site. The results strongly support the proposal of Bailey and Hawksworth to regard P. discolor as a distinct species.     

Monoterpenoid and morphological differentiation within Pinus cembroides

Wood of Pinus cembroides from 232 trees sampled in 23 localities throughout its range in the United States and Mexico has been analysed for monoterpenes. With a few exceptions the trees of the continental populations from the state of Hidalgo northward produced α-pinene as the main monoterpene. The majority of trees of the southern populations (subsp. orizabensis) from the state of Puebla and adjoining regions of the neighboring Veracruz and Tlaxcala produced 3-carene in appreciable amounts. The population from Baja California (subsp. lagunae) produced monoterpenes of the sabinene family (thujene, sabinene, γ-terpinene, terpinolene and $\text{p-}\text{cymene}$) as the main constituents. The variations in morphological characters, including number of needles per fascicle, number of stomatal rows in needles, number of resin canals in needles, as well as other characters of the internal needle morphology, needle length and thickness, needle retention, thickness of seed shells, bark and cone characteristics, rate of growth and cotyledon number either paralleled or did not contradict the chemical variations. The evolution of the southern populations (subsp. orizabensis) into a separate group most likely resulted from geographic separation and climatic differences, while the evolutionary separation of the Baja California trees (subsp. lagunae) was connected with northwestern movement of the California coastal region from Miocene onward and the formation of Sierra Madre Occidental resulting in separation of this population from the mainland populations by the gap of the Gulf of California and the coastal mountain ranges.     

Mountain Pine Beetles Use Volatile Cues to Locate Host Limber Pine and Avoid Non-Host Great Basin Bristlecone Pine

The tree-killing mountain pine beetle (Dendroctonus ponderosae Hopkins) is an important disturbance agent of western North American forests and recent outbreaks have affected tens of millions of hectares of trees. Most western North American pines (Pinus spp.) are hosts and are successfully attacked by mountain pine beetles whereas a handful of pine species are not suitable hosts and are rarely attacked. How pioneering females locate host trees is not well understood, with prevailing theory involving random landings and/or visual cues. Here we show that female mountain pine beetles orient toward volatile organic compounds (VOCs) from host limber pine (Pinus flexilis James) and away from VOCs of non-host Great Basin bristlecone pine (Pinus longaeva Bailey) in a Y-tube olfactometer. When presented with VOCs of both trees, females overwhelmingly choose limber pine over Great Basin bristlecone pine. Analysis of VOCs collected from co-occurring limber and Great Basin bristlecone pine trees revealed only a few quantitative differences. Noticeable differences included the monoterpenes 3-carene and D-limonene which were produced in greater amounts by host limber pine. We found no evidence that 3-carene is important for beetles when selecting trees, it was not attractive alone and its addition to Great Basin bristlecone pine VOCs did not alter female selection. However, addition of D-limonene to Great Basin bristlecone pine VOCs disrupted the ability of beetles to distinguish between tree species. When presented alone, D-limonene did not affect behavior, suggesting that the response is mediated by multiple compounds. A better understanding of host selection by mountain pine beetles could improve strategies for managing this important forest insect. Moreover, elucidating how Great Basin bristlecone pine escapes attack by mountain pine beetles could provide insight into mechanisms underlying the incredible longevity of this tree species.     

Terpenoid and morphological variability of Pinus quadrifolia and its natural hybridization with Pinus monophylla in Northern Baja California and adjoining United States

Mono- and sesquiterpenoids from wood of P. quadrifolia and that of the southwestern form of P. monophylla were analysed by GLC. Concurrently the number of needles per fascicle, the number of resin canals, and the number of abaxial and adaxial stomatal rows towards the tip, in the middle, and towards the base of the needles were determined. The data covered 90 trees from 8 pure populations from southernmost California and northern Baja California and 21 trees from a mixed stand in Baja California.α-Pinene, mycene, α-amorphene, γ-muurolene, germacrene D, and to a very minor extent camphene, β-pinene, limonene, β-phellandrene and β-ylangene percentages could be used for species identification. The number of needles per fascicle, the number of resin canals, and the number of abaxial stomatal rows afforded appreciably sharper identification, however. Statistical treatment of chemical and morphological data from pure stands did not give evidence for introgression of the two species. A definite intermediacy, rather limited in extent, could be demonstrated for the mixed stand, however.     

Volatile terpenoids from aeciospores of Cronartium fusiforme

Aeciospores of Cronartium fusiforme isolated from slash pine (Pinus elliottii) trees were analyzed for volatile terpenoids by GLC and GLC-MS. α-Pinene, β-pinene, Δ³-carene, myrcene, linonene, β-phellandrene, and δ-terpinene were the major monoterpenoid hydrocarbons present with only traces of camphene. A number of monoterpenoid alcohols were also present of which terpinen- 4-ol predominated. Among the various acyclic sesquiterpenes present, β-farnesene and β-citronellol were identified. Several aromatic compounds were also observed, including o-cresol.     

The variable climate response of Rocky Mountain bristlecone pine (Pinus aristata Engelm.)

Recent increases in temperature over the semi-arid western United States have been shown to exacerbate drought, reducing streamflow, and increasing stress on ecosystems. Our understanding of the role temperature played during drought in the more distant past is far from complete. While numerous tree-ring proxy records of moisture provide evidence for past extreme droughts in this region, few contemporaneous tree-ring proxy records of temperatures exist. This limits our ability to evaluate the variable influence of temperature on drought over past centuries and to contextualize the present interplay of moisture and temperature during more recent drought events. It is also important to understand the complexity of climatic interactions that produced drought under natural variability prior to evaluating the potential impacts of future climate change. In response to this knowledge gap, we undertook the first extensive evaluation of climate sensitivity in Rocky Mountain bristlecone pine (Pinus aristata Engelm.), focusing on the potential for developing new multi-century proxy records of both temperature and precipitation. We isolated dominant patterns of growth variability among trees from ten ring-width datasets across the Southern Rocky Mountains of Colorado and New Mexico and assessed their response to climate. We utilized both an empirical orthogonal function (EOF) analysis and a modified form of hierarchical cluster analysis to produce time series representing growth patterns in P. aristata. The results indicate a widespread June drought stress signal with a high potential for multi-millennial reconstruction. We also found a positive minimum temperature response during late summer, evident only at lower frequency and co-occurring at locations with the June drought stress signal. The potential for temperature reconstruction will require further investigation into the physiological linkages between P. aristata and climate variability. The presence of multiple climate responses within P. aristata sampling sites highlights the need for particular care when including P. aristata in regional climate reconstructions.     

灵空山油松-辽东栎林乔木树种群落学特征及空间分布格局

参照CTFS技术规范对灵空山自然保护区油松(Pinus tabuliformis)-辽东栎(Quercus wutaishanica)林的乔木进行每木调查的基础上分析了不同乔木树种的群落学特征及空间分布格局。结果表明,4hm2样地中共有乔木树种25种,总计8210株。隶属于12科18属,种的区系以东亚、温带亚洲及中国特有分布类型为主。油松、辽东栎是群落中的共优种,个体占总个体的86.81%,杜梨(Pyrus betulifolia)、白蜡树(Fraxinus chinensis)、漆树(Toxicodendron vernicifluum)为群落中主要的伴生种。优势种的径级结构与所有乔木个体的径级结构相似,近似于倒"J"形,以小径级个体居多,林下更新良好。主要树种的空间分布格局在小尺度上表现为聚集分布。油松小径级个体在20 m以内的尺度上呈聚集分布,当空间尺度超过60 m时表现为随机分布,大径级个体在所有尺度上均呈随机分布;辽东栎不同径级个体在小尺度范围内呈现聚集分布,随着尺度增大由随机分布趋于均匀分布。主要树种除漆树外在坡度较缓的生境下分布密度大。灵空山自然保护区油松-辽东栎林中油松种群优势地位更显著,各个乔木种群处于相对稳定状态。     

Allozyme diversity in Pinus virginiana (Pinaceae): intraspecific and interspecific comparisons

Two of the four members of subsection Contortae of the genus Pinus occur in the southeastern United States: Pinus virginiana, which ranges throughout the southern and central Appalachian Mountains, and P. clausa, which is restricted to Florida and southern Alabama. We examined allozyme variation within P. virginiana and genetic relationships between this species and the two varieties of P. clausa (var. clausa and var. immuginata). P. virginiana maintains more genetic diversity at both the species (Hes = 0.139) and population (Hep = 0.128) levels than the other three species in the subsection, which may reflect the combination of its widespread distribution and the absence of cone serotiny. Genetic differentiation among populations in P. virginiana was relatively low (GST = 0.053), but significant contrasts in allozyme frequencies and genetic diversity were apparent between populations to the northwest vs. outheast of the Appalachian Mountains. These regional differences likely resulted initially from historical processes that occurred during the Pleistocene and early Holocene, and have been reinforced by modern selective pressures and barriers to gene flow. The mean genetic distance between populations of P. virginiana and P. clausa (D = 0.071) was greater than that between populations of the two varieties of P. clausa (D = 0.012), which suggests that the two varieties diverged at some point after the separation of the two species.     

Differences between Pinus pinea and Pinus pinaster as bioindicators of polycyclic aromatic hydrocarbons

The potential of Univariate and Multivariate Analysis and specifically Principal Components Analysis (PCA), has been employed to assess the performance of pine needles as bioindicators of polycyclic aromatic hydrocarbons (PAHs) and particularly emphasize the crucial importance of making a distinction between different pine species if more than one is sampled. Four sampling sessions were done in 29 sites and needles of two common pine species (Pinus pinaster Ait. in 19 sites and Pinus pinea L. in 12) were collected and analysed using gas chromatography-mass spectrometry (GC-MS) and PCA. The results obtained indicated significant differences between species, attributed to their different morphology. The mean total PAH concentration of the P. pinaster needles are over two times higher than P. pinea's. This difference is lower when the results are presented in lipid weight, but still statistically significant. Samples from the two sites with adjacent trees reinforce these conclusions, showing significant differences in terms of PAH mean concentration and aromatic ring patterns.     

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.     

Fossil records of subsection Pinus (genus Pinus,Pinaceae) from the Cenozoic in Japan

Extant pines of subsection Pinus (section Pinus, genus Pinus, Pinaceae) are predominantly distributed in Eastern Asia. However, the extent of diversification in the section has yet to be fully clarified. We reviewed fossil records of subsection Pinus from Japan and collected permineralized materials, in which anatomical details are preserved for better understanding of the diversification. Our results suggest that this subsection appeared in Japan no earlier than the Middle Eocene, with extant species (i.e., Pinus densiflora and Pinus thunbergii) appearing around the beginning of the Pleistocene. Pinus fujiii (Early Miocene to Early Pleistocene) is inferred to have a close affinity to P. thunbergii based on the medial arrangement of its leaf resin canals. Additionally, P. fujiii has a similar cone morphology to those of extant species living in China, bridging the morphological gap between P. thunbergii and Chinese relatives of P. thunbergii as inferred by molecular phylogenetic analyses. Our results also suggest that taxonomic revisions of Pinus miocenica and Pinus oligolepis are required among the Japanese fossil species reported to date.     

Monoterpenoid differentiation in relation to the morphology of Pinus remota

Wood of Pinus remota from 126 trees sampled in 13 localities throughout of its ranges in the United States and Mexico has been analysed for monoterpenes. The trees of P. remota produced mainly α-pinene and limonene with ranges for population means of 68.0–92.3% and 3.2–16.9%, respectively. Limonene was useful for distinguishing this species from subsp. cembroides. Small chemical differences were found between the eastern and western variants of P. remota. The chemical differences between the trees of P. remota from the United States and the trees of the newly described species, P. catarinae, from Mpo Santa Catarina near Saltillo, Mexico, were also very few and quantitatively small. The variations in morphological characteristics paralleled the chemical differentiation. The chemical and morphological data substantiated the species status of P. remota, and suggested the status of a mere marginal variant for P. catarinae.     

The role of anthropogenic influences in a mixed-conifer forest mortality episode

Abstract. Anthropogenic influences on recent tree mortality in mid-montane mixed conifer forests of southern California, USA, and northern Baja California, Mexico, were investigated. The Pinus jeffreyi-Abies concolor phase of the mixed-conifer montane community was sampled at three sites, characterized by: (1) low levels of air pollution and long-term fire suppression (Holcomb Valley in the San Bernardino Mountains, California); (2) chronic, severe air pollution and long-term fire suppression (Barton Flats in the San Bernardino Mountains, California); and (3) little air pollution and no fire suppression (La Corona Arriba in the Sierra San Pedro Martir in Baja California, Mexico). Similar percentages of dead standing trees, around 14 %, were found at both San Bernardino sites, but a comparatively low level, 4 %, was found in the Mexican forest. Anthropogenic effects, in particular fire suppression, may play an important role in enhancing the impact of natural stresses on the dynamics of mixed conifer forests.     

EFFECTS OF TREE AGE ON SECONDARY XYLEM AND PHLOEM ANATOMY IN STEMS OF GREAT BASIN BRISTLECONE PINE (PINUS LONGAEVA)

Xylem and phloem tissue samples were collected from various-aged Great Basin bristlecone pine (Pinus longaeva D. K. Bailey) stems in southern Utah and southeastern California to determine whether the vascular cambia of older trees produce fewer xylem rays, shorter-lived xylem and phloem ray cells, fewer phloem sieve cells, and a thinner phloem. Increment cores were examined to determine whether ‘aged’ cambia produced narrower tracheids that might reduce water translocation. Sapwood thickness was measured and sapwood growth layers were counted on these cores. Regression and Classification and Regression Tree (CART) analyses of sample data found no age-related changes in cambial products. Phloem and xylem production appeared normal at all ages, with no evidence of cambial malfunction.     

Longevity of needle fascicles of Pinus longaeva (Bristlecone pine) and other North American pines

Summary Variation in leaf longevity of gymnosperms has received surprisingly little attention despite its likely adaptive significance. Pinus longaeva, a pine of arid, subalpine environments in the western United States, has the record among conifers for needle longevity, with a maximum dwarf shoot (needle fascicle) retention time of up to about 45 years. Most low elevation pines have dwarf shoot retention times (DSRs) of two to four years. Literature data for the 37 species of Pinus native to the United States and Canada and field data for eight taxa (21 populations) of pines growing at various elevations in California each show a strong positive correlation between elevation and DSR, respectively, r=+0.65, df=35, p<0.001 and r=+0.82; df=19, p<0.001. We propose that extended needle fascicle longevity represents an adaptation to arid and especially high elevation environments. Field data from native stands and common gardens indicate that differences between taxa in DSR relate to both genetic and environmental factors. When grown at the same sites certain species (eg, P. longaeva, P. monophylla) had much longer DSRs than others, indicating a genetic basis for differences in needle fascicle longevity. For six of seven taxa that were each studied at more than one elevation there was a statistically significant increase in DSR in going from the lowest to the highest elevation site, indicating strong environmental control of needle fascicle longevity.The physiological control of dwarf shoot senescence and abscission is poorly understood. For P. longaeva dwarf shoots of a particular age class are not shed simultaneously; rather there is a more or less gradual attrition of dwarf shoots from the long shoot. Although different types of long shoots of pines are known to differ physiologically, for P. longaeva there was no consistent difference in DSR on various types of lateral long shoots (eg, vegetative, pollen cone-bearing, seed cone-bearing), nor was there a statistically significant difference in DSR on trunks versus on their lateral long shoots. In addition, for P. contorta ssp. bolanderi and P. muricata needle fascicle longevity was not affected by the degree of edaphically induced dwarfing (ie, stunting) of the trees.     

秦岭中西部油松径向生长对气候因子的响应差异研究

在气候变暖背景下,树木径向生长对气候因子的响应会随区域干湿变化而有所差异。秦岭属于气候敏感区和生态脆弱区,南北气候特征差异明显,分析气候变化背景下树木径向生长对气候因子及干旱事件的响应,对准确预测未来气候变化对树木生长的影响至关重要。为探究该地区不同干湿环境下油松对气候因子及干旱事件响应的特点及差异,共采集秦岭中西部南北坡共4个样点的油松树轮样芯,利用树木年轮生态学的方法,分析各地油松年表与气候因子之间的关系,通过计算抵抗力、恢复力、恢复弹力等指标探究树木径向生长对干旱事件的应对能力,结果表明：1)在西部和北坡的3个样点,油松径向生长主要与前一年7—9月、当年5—7月的气温呈显著负相关,与当年5月降水呈显著正相关,在中部南坡油松径向生长主要与当年2—4月、9月气温呈显著正相关,与当年4月降水呈显著负相关;2)生长季气温升高所引发的水分亏缺,是研究区西部和北坡油松径向生长受限制的主要原因,且中部南坡油松生长受气温和干旱因子的制约要明显弱于其它3个样点;3)西部北坡油松对干旱的抵抗能力、恢复能力及恢复弹力均弱于西部南坡及中部北坡油松。在未来对研究区树木的管理与保护工作中,应更加关注西部北坡...