Health of North American Forests L’état de santé des forêts nord-américaines Sanidad de los Bosques de América del Norte Produced by the Atmospheric Changes and Forests Study Group of the North American Forestry Commission in cooperation with: Établi en collaboration par le Groupe d’étude sur les changements atmosphériques et les forêts de la Commission forestière pour l’Amérique du Nord : Informe producido por el Grupo de Estudio de Cambios Atmosféricos y Bosques de la Comisión Forestal de América del Norte, en cooperación con: Canada/Canadá J. Peter Hall Laszlo Magasi Lester Carlson United States/États-Unis/Estados Unidos Ken Stolte Elvia Niebla Mexico/Mexique/México Maria de Lourdes de la I. de Bauer Carlos E. Gonzalez-Vicente Tomás Hernández-Tejeda North American Forestry Commission Report/ Rapport de la North American Forestry Commission/ Informe de la Comisión Forestal de América del Norte Published by / Publié par / Publicado por Canadian Forest Service / Service canadien des forêts Science and Sustainable Development Directorate/ Direction des sciences et du développement durable Ottawa, 1996 Cover: Aspen in an alpine meadow. (CFS—Petawawa National Forestry Institute) Couverture : Peuplier dans un pré alpestre. (SCF—Institut forestier national de Petawawa) Tapa: Álamo temblón en un prado alpestre. (CFS—Petawawa National Forestry Institute) Contents / Table des matières / Indice Health of North American Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 North American Biomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Inside back cover L’état de santé des forêts nord-américaines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Biomes nord-américains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 3 de la couverture Sanidad de los Bosques de América del Norte . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Biomasas en América del Norte . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reverso de la contratapa Appendices/Annexes/Anexos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 © Her Majesty the Queen in Right of Canada 1996 Catalogue No. Fo42-252/1996 ISBN 0-662-62141-7 Copies of this publication may be obtained free of charge from: Natural Resources Canada Canadian Forest Service Ottawa, Ontario K1A 0E4 A microfiche edition or photocopies of this publication may be purchased from: Micromedia Ltd. 240 Catherine St., Suite 305 Ottawa, Ontario K2P 2G8 Editing and Production: Paula Irving Design and Layout: Danielle Monette Canadian Cataloguing in Publication Data Main entry under title: Health of North American forests = L’état de santé des forêts nord-américaines = Sanidad de los bosques de América del Norte (North American Forestry Commission Report) Text in English, French and Spanish. Includes bibliographical references. ISBN 0-662-62141-7 Cat. no. Fo42-252/1996 1. Forest health—North America. 2. Sustainable forestry—North America. 3. Forest conservation—North America. 4. Forest management—North America. I. Hall, J. Peter. II. Canadian Forest Service. Science and Sustainable Development Directorate. III. North American Forestry Commission. IV. Title: L’état de santé des forêts nord-américaines. V. Title: Sanidad de los bosques de América del Norte. VI. Series. SD387.S87H42 1995 Printed on recycled paper 634.9’6 C95-980298-3E PRINTED IN CANADA Printed on alkaline permanent paper Health of North American Forests Contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Forest Biomes in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Pollution and Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Acid Deposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Ground Level Ozone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Nitrate Pollution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Stress in Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Forest Harvesting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Forest Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Pest Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Assessing Forest Health . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Health of North American Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 1. Temperate Needleleaf Forests or Woodlands. . . . . . . . . . . . . . . . 11 2. Temperate Broadleaf Forests or Woodlands . . . . . . . . . . . . . . . . 12 3. Mixed Mountain and Highland Systems . . . . . . . . . . . . . . . . . . . 13 4. Subtropical and Temperate Rain Forests . . . . . . . . . . . . . . . . . . . 14 5. Evergreen Sclerophyllous Forests . . . . . . . . . . . . . . . . . . . . . . . . . 14 6. Tropical Dry Deciduous Forests or Woodlands . . . . . . . . . . . . . . 15 7. Tropical Humid Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Selected Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Appendix I. Major Tree Species in Each Biome . . . . . . . . . . . . . . . . 53 Appendix II. Damaging Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Appendix III. Parasitic Plants and Hosts . . . . . . . . . . . . . . . . . . . . . . 64 6 Health of North American Forests Abstract Forests are environmentally and economically important in North America, and forest health is monitored to enable policy decisions to be made on factors affecting forest health. Forests are subject to several stresses, natural and human, and these stresses have various effects. Data for monitoring systems are used to separate the causes of observed damage to forests. North American forests are divided into seven biomes and forest health is described for each. Mortality in North American forests is generally within the normal range for forests and is mostly caused by natural thinning. A variety of known insects, diseases, and abiotic factors killed trees and damaged tree crowns. Some forests are stressed by land use practices. Incidence of pollution-like symptoms occurred in several biomes and on many species. Where these symptoms were observed, trees are being studied to determine the cause and extent of damage. Introduction This publication is a result of a cooperative effort by the Atmospheric Changes and Forests Study Group of the North American Forestry Commission (NAFC). The report is an overview of the state of the health of North American forests based on information from Canada, the United States, and Mexico. The forests described range from the boreal conifers to the tropical hardwoods, and from coastal rain forests to mountain timberline forests. These forest ecosystems are a source of clean air and water and are areas for homes for many of our citizens. They provide recreational opportunities and a habitat for wildlife, and yield pulp and paper, lumber, food, and firewood. Consequently, North Americans are very concerned about the health of these ecosystems. The sustainable development of these forests is essential to our economic, biological, and spiritual health. Forests are subject to natural and anthropogenic stresses that act singly or collectively to affect forest health. There is no universally accepted definition of forest health, except for poor health, which is evident when forest ecosystems are deteriorating or declining. When forests are damaged by insects, diseases, fire, or climatic events, they are probably only temporarily unhealthy because they are adapted to these stresses and they can recover from them. Stresses such as air pollution or potential climate change may permanently damage forest health, because forests have not adapted to these additional stresses. Anthropogenic air pollution as a primary or contributing source of stress is present to varying degrees in most North American forest ecosystems. Pollution has been affecting European forests for many years. In Canada, air pollution has damaged conifer forests in British Columbia and Ontario, and birch forests near the eastern coast. In the United States, the red spruce forests in the Appalachian Mountains and ponderosa pine in the San Bernardino and Sierra Nevada mountains have been affected. The pine and fir forests in the valley of Mexico have also been damaged by atmospheric pollutants. Forests in all three countries remain potentially at risk from atmospheric pollution. To protect our forests, we must know the state of their health. This report describes the mortality and crown condition of trees in these forests as affected by various stresses. The Atmospheric Changes and Forests Study Group expects to report on the state of the health of North American forests regularly. Forest Biomes in North America The health of forests is discussed in seven forested biomes or natural regions as described for the Man and Biosphere Program (see the map on the inside back cover). The forest types, major species, and forest stresses are described briefly for each biome. 1. Temperate Needleleaf Forests or Woodlands This biome dominates the northern latitudes from the Arctic tree line south to the Great Lakes and the hardwood forests of the eastern United States. Forests in this biome consist of the boreal and mixedwood species with a variety of conifers and hardwoods. The major species include white spruce, black spruce, tamarack, balsam fir, eastern hemlock, white pine, jack pine, lodgepole pine, sugar maple, white birch, yellow birch, aspen, and balsam poplar. Many of these ecosystems are dominated by wildfires, and the regeneration of black spruce and jack pine depends largely on wildfire. These forests have been harvested for many years and large clearcuts predominate in the north. In the south, extensive areas have been converted to agricultural use. 2. Temperate Broadleaf Forests or Woodlands This biome is composed primarily of hardwood species and a mixture of conifers dominates in the southeast. It includes forests in southern Canada and in the United States east of the Prairies. It is bordered on the north by the temperate needleleaf forests and in the east and south by the Atlantic Ocean and the Gulf of Mexico. The major broadleaf species include beech, yellow poplar, hickory, yellow birch, black cherry, sugar maple, red maple, and oaks. The major conifers include white pine, eastern hemlock, red spruce, and balsam fir. In the southern Health of North American Forests part of the biome, loblolly pine, slash pine, shortleaf pine, white oak, and scarlet oak predominate. This is a species-rich biome with many forest types. Much of the original forest was cleared for settlement after 1600. During the last century, much of the agricultural land has reverted to forest. The forests are now affected by a wide variety of insects, diseases, and pollutants. 3. Mixed Mountain and Highland Systems These are continental interior forests ranging from latitude 15 to 60°N on sharply changing topographical relief. Typically these forests begin about 2000 m above sea level and go up to about 3500 m in the southwest, and begin near sea level farther north and go up to around 3000 m. Dominant species in the northern part of the biome include Engelmann spruce, alpine fir, lodgepole pine, Douglas-fir, subalpine fir, ponderosa pine, white fir, red fir, trembling aspen, and oaks. In the south, the species composition changes to a rich mixture of species. So far, 128 different species of oaks have been identified in Mexico, and there may be more species to be described and classified. Similarly, 54 different species of pines are known in these forests, along with 9 species of true firs, 4 species of Douglas-fir, 2 species of spruce, and 3 species of cypress. Many of these ecosystems are fire-dominated, with additional localized harvesting and land clearing activities. The forests are affected by insects, diseases, and parasitic plants. Some species are affected by pollutants near Mexico City, Jalisco (Guadalajara), and Nuevo Leon (Monterrey). 4. Subtropical and Temperate Rain Forests This biome consists of the Pacific coastal forests from Alaska to California and is dominated by western red cedar, western hemlock, Sitka spruce, Douglas-fir, Amabilis fir, yellow cypress, mountain hemlock, alpine fir, subalpine fir, redwoods, ponderosa pine, white fir, and red fir. These forests are valuable and have been extensively harvested. Insect defoliators, bark beetles, and stem and root diseases also affect these forests. Forest wildfires are not common, but forest management activities have been quite intensive. 5. Evergreen Sclerophyllous Forests Forests in this biome are characterized generally as dryland forests. Typically the species are small and have sclerophyllous (leathery) evergreen needles or leaves. Growth rates are typically slow because of low rainfall and the low fertility of the soils. The dominant tree species are pinyon, juniper, oak, manzanita, ceanothus, and chamise. These forests occupy higher 7 elevations in desert areas and lower elevations and warmer slopes of mountains in the more southern latitudes. Some of these forest communities are well adapted to frequent wildfires which maintain these species in the biome. 6. Tropical Dry Deciduous Forests or Woodlands This biome occurs in western and southern Mexico and in the Yucatan Peninsula. Again there is a rich species mix with 27 different species of Acacia; A. coulteri and A. farnesiana are typical. Other species include Alvaradoa amorphoides, Bursera spp., Ceiba acuminata, C. aesculifolia, Lysiloma divaricata, Prosopis laevigata, Fraxinus purpusili, Piscidia communis, Lysiloma bahamensis, Pithecellobium dulce, Cyrtocarpa procera, Cordia dodecandra, Tabebuia chysantha, and Gliricidia sepium. These forests grow from 0 to 1900 m, and the annual mean temperature is 20–29°C. There is one dry season (5–8 months) and one rainy season. Most forests are on rocky soils and hillsides of low fertility. Some of these species have a high economic value. The main problems for these forests are deforestation, wildfire, and grazing. 7. Tropical Humid Forests This biome comprises the coastal forests in Mexico from Veracruz to the southern Yucatan Peninsula. The dominant tree species are Dialium guianense, Terminalia amazonia, Swietenia macrophylla, Cedrela mexicana, Vochysia hondurensis, Andira galeottiana, Swartzia panamensis, Calophyllum brasiliense, Guatteria anomala, Aspidosperma megalocarbon, Manilkara zapota, Bursera simaruba, Dendropanax arboreus, Tabebuia rosea, Alchornea latifolia, Guarea excelsa, Celtis monoica, Brosimum alicastrum, Pimenta dioica, Platymiscium yucatanum, Robinsonella mirandae, Astronium graveolens, Zanthoxylum kellermanii, Ceiba pentandra, Sabal mexicana, Trichilia minutiflora, Bucida buceras, Metopium brownei, Terminalia oblonga, Virola guatemalensis, Zuelania guidonia, Sterculia mexicana, Barnoullia flammea, Blepharididium mexicanum, Pseudolmedia oxyphyllaria, Ficus insipida, Bravaisia integerrima, and Quararibea funebris. There are many epiphytic plants of the Bromeliaceae and Orchidaceae families. In some cases, there are 34 different species of plants in 2000 m2 and 735 trees/ha. These forests grow in flat lands, and deep, rich, well-drained soils. The species mentioned grow from sea level up to 1000 m. The average precipitation is from 1500 to 3000 mm, and the annual mean temperature is above 29°C. These forests have been extensively clearcut for agriculture and grazing. There is shift cultivation practiced in which the trees are cut and burned, and the soil is cultivated for a few years then abandoned. After that, the vegetation regrows and 8 Health of North American Forests the cycle begins again. Land use change is thus a major factor affecting the health of these forests. Wildfires are common in these forests. Sometimes the clearcut areas are used as grasslands. Erosion in areas without vegetation can be severe due to the heavy rains. Some damaging insects and diseases are present, and damage on plants by pollutants emitted by oil refineries is common. Pollution and Forests Air pollution was first recognized in the nineteenth century as potentially harmful. Because natural ecosystems have a built-in resilience to stress, it took many years for air pollutionrelated changes in biological organisms to become obvious. However, the science community now accepts that air pollution affects forests, lakes, agricultural crops, cultural monuments, and human health. Air pollution from individual point sources will kill trees and destroy complete ecosystems typically somewhere within 25–50 km of the source. In North America, considerable damage was done to forests from point sources during the nineteenth and early twentieth century. More recently, concerns have arisen regarding the possible effects of both point-source and regional pollution in decline and death of trees and forest ecosystems. Damage caused by regional transport of pollutants is difficult to detect, and forest declines in areas of low and high levels of acid deposition have occurred. The effects of pollution damage have been frequently duplicated under laboratory conditions; however, under ambient levels of pollution, the mechanism of cause and effect is not clear and has seldom been demonstrated. Removal of the stress by reduction of emissions, however, usually results in the recovery of the affected vegetation, suggesting a cause–effect relationship. Many pollutants are affecting forests in North America; acidic deposition, ozone, nitrates, various particles, and heavy metals are best known. Transported pollutants affecting forests: Gaseous Sulfur compounds Nitrogen compounds Photo-oxidants Hydrocarbons Pollutant mixtures Particulates Acidic (S and N) compounds Heavy metals Hydrocarbons Solutions Oxidants Acid rain/fog Hydrocarbons Direct/indirect effects of air pollution: Direct effects 1. 2. 3. 4. 5. 6. 7. nutrient leaching from foliage ozone damage and decreased photosynthetic efficiency ozone damage and increased foliar leaching sulfur dioxide damage foliar fertilization and increased winter injury foliar fertilization and altered nutrient allocation growth altering substances Indirect effects 8. nutrient leaching from soil 9. aluminum mobility and toxicity 10. heavy metal toxicity Concerns have also recently been raised about volatile organic compounds (VOC) and UV-B radiation. These pollutants originate from various anthropogenic sources and are transported over long distances and over extensive areas. Acid Deposition Acid deposition consists of wet and dry deposition of SOx and NOx. Acid deposition/acid rain affects vegetation by interacting with the leaf surface. It affects soils by inhibiting nutrient uptake by trees (nitrogen/phosphorus) and releasing toxic (aluminum) elements. Acidic deposition levels of 20 km • ha-1 • yr-1 or greater adversely affect lakes and streams in North America. Biomes 1 and 2 are exposed to pollutants in this way. Air pollution is an additional stress on North American forests beyond the normal damaging factors. These added stresses can predispose forest trees to damage from other insects and diseases that otherwise might cause less damage. Emissions of acidic pollutants may have peaked and declined slightly over much of North America; however, in all areas pollutants are still accumulating in soils, water, and vegetation. In some areas emissions of pollutants are still increasing. Ground Level Ozone Tropospheric (ground level) ozone is produced in the earth’s lower atmosphere from photochemical reactions involving oxides of nitrogen and volatile organic compounds (NOx/VOC) in the presence of sunlight. Ozone damages plants by entering through the open stomates and affects mesophyll cells in the leaf. It builds up in the lower atmosphere especially on hot days in cities and flows out into the surrounding countryside. Health of North American Forests Thresholds for damage to vegetation is 50–80 parts per billion. Agricultural leaf crops are widely affected causing considerable economic losses annually. This tropospheric ozone is increasing at a rate of 1–2% annually. Ozone air pollution in the temperate broadleaf forests of biome 2 is of concern. Ozone levels are highest in the midAtlantic and upper southeastern coastal plain areas. The effects are magnified in the southeastern part of the biome. Plants are exposed longer because they flush early, and humidity levels are higher, facilitating the uptake of ozone. Ozone-sensitive tree species such as black cherry, sassafras, and yellow poplar are injured in both the upper and middle of the crowns in more than 60% of trees. High ozone exposures cause ozone injury when relative humidity and soil moisture are high. During periods of drought, damage is limited because plant stomates close. In biomes 2 and 3, the temperate broadleaf and mixed mountain and highland biomes, trees are frequently exposed to phytotoxic concentrations of ozone. Ozone stress typically occurs in the mixed-coniferous forests at higher elevations (>1500 m). In the San Bernardino Mountains of southern California, and the southern and central Sierra Nevada Mountains farther north, ozone is particularly injurious to ponderosa pine, Jeffrey pine, and to a lesser extent to black oak. Damaged foliage, loss of needles, and growth reductions have occurred. A similar situation to the one described for the San Bernardino Mountains has been detected in southern mountainous areas of Mexico City where at altitudes of 2850–3500 m, Pinus hartwegii, one of the nine native species to the Valley of Mexico, was highly sensitive to ozone. During the last 3 years, the O 3 monitoring data for the ozone have shown values that almost daily exceed the Air Quality Standards and very frequently remain for several hours a day, well above the threshold for damage to vegetation. The observed damage on pines first at Ajusco and later at the national park Desierto de los Leones located farther southwest of Mexico City is severe and the population of P. hartwegii in certain areas is declining or dying. The patterns of growth of P. hartwegii using standard dendrochronological techniques show a marked decline in growth since the early 1970s at both sites (Ajusco and Desierto de los Leones) and there are no signs of subsequent recovery. In terms of typical oxidantinduced symptoms, there is a variability in the response of the pine species in the area and among individuals within species. Nitrate Pollution The source of nitrate pollution includes automobile, agricultural, and industrial emissions. There are increasing concerns about nitrate deposition because its levels are increasing at the same time as sulfate levels are decreasing in many areas. 9 Continued deposition of nitrogen will increase the amount of nitrogen in forest ecosystems, and depending on the nature of the soil will increase the capacity for nitrification and the movement of nitrogen to surface water. Vegetation growth can be expected to increase where nitrogen is a limiting factor, and where the deposition is not accompanied by elements interfering with nutrient uptake and/or where it introduces additional stress. Continued nitrogen deposition affects other components of the ecosystem including mycorrhizae, insects, pathogens, and wildlife. Present ecosystem interactions may be changed in unpredictable ways. Nitrogen deposition is of particular concern in biome 2 and the southern parts of biome 1. In biome 5, evergreen sclerophyllous forests, nitrate emissions (NOx) are greatest near major cities. While sulfate emissions (SOx) are generally decreasing, nitrate emissions have been steadily increasing over the last century. The effect of increased deposition of nitrates is to fertilize trees. When this happens the trees do not harden off properly for their dormant season. This can result in extensive foliar damage from low temperatures or droughts because the trees are not able to respond to their changed environment. The factors associated with mortality in the past have now been supplemented by pollutants of various types. The pollutants therefore are causing additional stress by making trees more susceptible to insects, diseases, climatic extremes, or nutrient stresses. Pollutants may also trigger a decline in the health of forests already stressed by other factors. Stress in Forests Stress is a constant and natural part of the forest ecosystem. Stress varies over time and space, and individual trees and forests react to it by tolerating and recovering, or by declining and dying. Chronic and acute stresses affect forest ecosystems. Chronic stresses influence forest ecosystems over long periods and over large areas. Examples include climate, soil parent material, topography, and competition. Acute stressors are more limited in space and time, and include insects, diseases, pollution, storms, and fire. Most natural stresses do not damage forest ecosystems beyond repair, and they recover to a state resembling the original conditions. Only in severe cases is an ecosystem driven over its threshold of resilience, in which case it acquires a new equilibrium, or it declines and degrades completely. Largescale forest declines are often thought to have been triggered by stress events (drought, frost, pollution) and may have been aggravated by a predisposing condition of many years of other stresses including pollutants. Natural competition causes stress and occurs when individual trees compete for growing space, solar radiation, water, 10 Health of North American Forests or essential nutrients. In natural forests more trees die of stress from competition than from any other factor. The “normal” mortality in unmanaged forests from natural thinning is about 1–3% annually. Other natural stresses may cause higher levels of mortality. Some of these are biological (animals, fungi, insects, nematodes, bacteria); others are climatic (drought, floods, storms). Wildfires cause extensive tree mortality but usually result in a new forest becoming established and are not considered harmful to forest health in the long term. Human (anthropogenic) stressors such as air pollution, harvesting, exotic forest pests, and forest management activities all add to natural stresses. Human-caused disturbances are common to forests that have been extensively exploited. Two instances of forest decline in eastern North America illustrate the interplay of natural and anthropogenic causes. In eastern Canada, a well-known decline and dieback sequence occurred in yellow birch beginning about 1925. Extensive areas of mortality occurred in disturbed and undisturbed stands and on various physiographic site types. Various insects and diseases were suggested as causes as were weather events, particularly droughts and unusually mild winters, but no cause and effect link was determined. Since approximately 1950, however, there has been a gradual recovery from the decline, and the species is now generally healthy. A case of an interaction between a foliar pathogen (Lophodermium sp.)and oxidant gases has been described as present in the mountainous forest “El Ajusco,” located south of Mexico City. Here Pinus hartwegii shows severe foliage damage in addition to the characteristic pollution-induced symptoms chlorotic banding and mottling of the leaves. Abundant fruiting bodies of Lophodermium sp. are present on the affected foliage. The magnitude of the damage is directly related to the age of the leaves and to the side of the tree exposed to the urban zone. In a high elevation forest park near Mexico City, the Desierto de los Leones, where the O3-induced symptoms on P. hartwegii are evident at certain altitudes and exposures, a massive forest decline of sacred fir was observed for the first time in 1982. It was similar to tree damage and decline described in Europe and North America. Trees located in the windward zone, exposed to air masses from Mexico City, were the most severely affected, especially trees at the distal ends of ravines. Within 2–3 years after the symptoms appeared, the trees were dead. Although there is much less information in relation to air pollution effects on Abies than on pines, it was concluded that the polluted air from Mexico City was an important factor in the decline of fir. However, drought, insects, mites, pathogens, and poor forest management have collectively contributed to this decline. The areas where the sacred fir trees died have been reforested with different pine species as well as with sacred fir. Most of those trees show already very clear oxidant- type symptoms and some of them are in poor health. Thus under those special conditions the zone can be considered a trial selection for tree tolerance to O3. Decline has occurred in red spruce forests over the past century in the Appalachian Mountains, and recent growth records suggest that it is far more widespread than previously. Severe droughts in the mid-1960s might have initiated the decline. However, since the decline occurred before the drought, and in nondrought areas, drought was probably not the primary cause. Recent research has indicated that a combination of site factors, climate, and sulfate/nitrate pollution are collectively causing the decline. Forest Harvesting Forest harvesting has occurred in much of the North American forests. Harvesting interrupts the natural succession in the forest, affects the water balance, causes soil erosion and compaction from harvesting equipment, and leaves logging slash which provides fuel for wildfires. Fires are suppressed to protect commercially valuable forests particularly in the temperate needleleaf forests. These actions have resulted in a successional preference for balsam fir rather than the fire-tolerant species jack pine and black spruce. The resulting balsam fir forests are prone to damage from defoliating insects such as the spruce budworm and the hemlock looper. All these factors result in areas of forest under increased stress. Harvesting of trees has often been done in ways that adversely affect forest health. Harvesting methods such as selective cutting and protection of wildlife habitat, old growth forests, streams, and soils all help avoid stress on forests. Forest Management Increases in the intensity of forest management may result in stress, particularly by conversion of stands from one species or species group to another. This may lead to an ecologically unsustainable forest and may at the same time be aesthetically displeasing. Examples include extensive conifer tree plantations in biomes 1 and 2 and in the southern parts of biome 4. Large monocultures are also often more susceptible to infestation by damaging insects and fungi than forests composed of a mixture of species. For example, second-generation stands of Norway spruce frequently have reduced growth rates compared with the previous forests on the same site. This is caused by lower pH resulting from spruce growing on soils that previously supported deciduous species. This causes nutrient stress on the new forests and makes them vulnerable to other stresses such as pollutants. Land was originally cleared for agriculture in North America, cultivated, and then reverted to forests as the land became Health of North American Forests impoverished. At present many of these lands support forests in the early successional stages and are very often under nutrient stress because many of the original nutrients were removed by other crops. Modern forest management emphasizes the ecological integrity of the forest over timber production. This approach is now being widely adopted in temperate and boreal forests, and criteria and indicators are being developed by councils directed by the United Nations Council on Economic Development. Pest Introduction There are several insects and diseases accidentally introduced to North America in the nineteenth century. They were not particularly damaging in their native habitat but caused extensive problems in their new environment because natural controls were lacking. Examples include chestnut blight, Dutch elm disease, beech bark disease, and balsam woolly adelgid. Both chestnut blight and Dutch elm disease have had such a devastating effect that the host species, American chestnut and American elm, have ceased to be a significant part of the Carolinian forest. These ecosystems have changed and now contain a different mixture of species. The balsam woolly adelgid has killed much of the higher elevation Fraser fir in eastern North America, leaving extensive areas of dead trees that are being replaced by other species. These stresses have had major effects on forest health. Forest ecosystems have responded by developing into different ecosystems; the original forest types have probably disappeared forever. 11 Current forest conditions and changes are measured annually. Tree mortality is evaluated; the rate considered normal for an unmanaged forest is 1–3%, caused primarily by thinning in natural stands. Higher levels may be caused by insects, disease, drought, frost, etc., singly or in combination. Forests that had normal or excess mortality from these causes would not be considered unhealthy. However, forests that had mortality from unknown causes or from other than normal causes for which forests are adapted would be considered in poor health. Tree growth may be used to evaluate the effects of pollutant stress on forests. In many instances it is difficult to detect changes in growth caused by pollution among the other factors affecting growth. Unless there is massive reduction in growth in polluted areas compared with growth in nonpolluted areas, it is difficult to use growth rates to detect pollution effects. Tree condition is evaluated for conifers and hardwoods. The types of damage from insects and diseases are described. The cause of defoliation and dieback is attributed to insects, diseases, or other factors. The symptoms of air pollution damage are easily confused with symptoms of damage from natural causes, and the correct identification depends on the experience of personnel trained to distinguish symptoms of air pollution damage from symptoms of abnormal climatic conditions, nutrient deficiencies, and the effects of insects and diseases. Symptoms are explained for both conifers and hardwoods. When damage not easily explicable by natural causes is observed, research is necessary to determine the cause of the damage. Health of North American Forests Assessing Forest Health To ensure forest sustainability, we first must know how our forests are functioning in an environment that has been to some extent changed by human activity. With this knowledge action can be taken to protect forests. These questions are asked during monitoring: Is this forest/tree different from the normal healthy forest/tree and if so what has caused the change? Are the changes caused by insects, diseases, drought, frost, or some unknown factor? Forest management agencies have established permanent sample plot systems in representative forest ecosystems, and results from these plots are synthesized for the major species in each of the seven biomes. Data collected are supplemented by the results of additional surveys conducted by national and local forest management organizations. There are several, localscale monitoring and research programs to determine the condition of the forest, measure annual changes, and evaluate trends in forest condition. The health of forest ecosystems is described for each biome. The assessment of forest health is based on tree mortality, tree condition, and pollution symptoms. The causes of mortality and crown damage are also described. 1. Temperate Needleleaf Forests or Woodlands The major stresses acting on this biome are various defoliating insects, root rots, and localized effects of harvesting and wildfire. Symptoms of air pollution have seldom been observed on trees in this biome. Tree Mortality Tree mortality among most species in this coniferdominated biome is mostly in the normal range of 1–3% for unmanaged forests. Causes of mortality were natural thinning, blowdown, and sporadic attacks by forest insects such as the spruce budworm, the balsam woolly adelgid, and the ragged spruce gall adelgid. 12 Health of North American Forests Normal levels of mortality also prevailed in the hardwoods, although damage levels were higher than in the conifers. Mortality of white birch, a species that had been stressed by severe foliage browning before 1986, returned to normal levels compared with previous years. Mortality in the aspen species was higher than normal and was caused by natural thinning, blowdown, drought, late spring frosts, defoliating insects, and root and stem rots. Tree Condition The conifers were generally in good condition. Local insect outbreaks or weather conditions affected some species. In particular, the spruce budworm, twig aphids, and mites caused defoliated spruces. Pine species were healthy and had no significant crown damage/defoliation. Some damage from canker and needle cast disease caused loss of needles on lodgepole pine, and a combination of insect pests, drought, and poor site quality affected the condition of red pine. A common rot, brown cubicle rot, also caused mortality in white spruce. Other damaging insects included the pitch nodule maker and a long-horned beetle; diseases included armillaria root rot and gall rust. Balsam fir, a host species for several defoliating insects, was quite healthy. The 50-year cycle of the eastern spruce budworm peaked in the 1980s and has since been declining. Some trees previously weakened by insects have died and there are localized infestations of the eastern spruce budworm and the hemlock looper. Eastern larch is occasionally infected with European larch canker causing gradual deterioration in crown condition, branch mortality, and occasional dead trees. As a result of cankers and other defoliating insects, decline of eastern larch is occurring in some regions, and mortality can be expected to increase above normal levels in the future. Birch crowns were affected to varying degrees by drought, storm damage, various insect pests (hemlock looper and the spearmarked black moth) and diseases, leaf spot, and root rots. Aspen crowns were damaged by insects, diseases, summer drought, and late spring frosts. Forest tent caterpillar and the large aspen tortrix have caused most of the crown damage. The crowns of largetooth aspen were generally healthy and had occasional marginal discoloration and chlorosis caused by drought. Pollution Symptoms In eastern Canada, severe foliage browning has led to the death of many northern white birch trees. The browning damage is related to the presence of acidic coastal fog which damages the foliage and causes branch and eventually tree mortality. Weakened trees die from this and other forest stresses such as armillaria root rot, having first been predisposed by the pollutants. In addition to the pollution damage on birch, occasional needle flecking and chlorosis were also observed on white pines and red spruce. These symptoms are consistent with those caused by ozone, but the actual cause has not yet been determined. 2. Temperate Broadleaf Forests or Woodlands The temperate broadleaf forest biome of the eastern United States encompasses a wide variety of forest ecosystems, of which oak–hickory is the major forest type extending from Texas in the south to the Dakotas in the north, and east to southern New England. These forests have been extensively stressed by damage from introduced pest species and their species composition has been altered during the past few decades. This forest type was formerly chestnut–oak, when American chestnut dominated these forests. Chestnut blight, introduced in 1900, has eliminated American chestnut, reducing it to an understory species seldom reaching tree size. Tree Mortality Mortality was low in these forests, in the normal range of 1–3%, and was caused by natural thinning as the stands mature and develop. Exceptions included higher than normal mortality in sugar maple caused by Armillaria sp. after the trees were stressed by other natural factors. Balsam woolly adelgid populations caused extensive mortality of Fraser fir in the southern Appalachians. Tree Condition The crowns of most conifer species were healthy. There was some defoliation and discoloration caused by the occasional presence of insects and diseases on the foliage. On hardwoods, there were occasional trace amounts of twig or branch dieback and marginal chlorosis or discoloration, and some damage from insects, diseases, and abiotic factors. American beech was healthy except for stem wounds or nectria canker, an introduced pathogen that severely damages beech. Red oak crowns were damaged slightly by pear thrips and localized drought stress. The crowns of sugar maple were mostly healthy. Trees are recovering from insect infestations and drought in various parts of the range. Symptoms of these stresses included discoloration and scorch of leaves and chlorosis, both symptoms consistent with drought. The major biotic stresses on this biome include gypsy moth, the southern pine beetle, eastern spruce budworm, balsam woolly adelgid, fusiform rust, and dogwood anthracnose. Gypsy moth is currently a major pest in these forests and continues to defoliate hardwoods over extensive areas and in isolated patches. It was introduced in 1869 and has since spread south and west, feeding mainly on oaks. Stressed trees are often attacked by two-lined chestnut borer and armillaria root rot. By Health of North American Forests 1991, gypsy moth had defoliated over 1.6 million ha and infested another 50 million ha. In the southern part of the biome, southern pine beetle continues to affect southern pines. More than 24 000 infestations occurred in 1991. Dogwood anthracnose occurs in the mountains and piedmont areas of several southern states and is spreading throughout these areas. The disease causes defoliation and mortality; in areas above 1000 m all trees may be killed. Oak decline is another major stress factor on the oak– hickory forest. It is a complex initiated by several factors: drought, site condition, foliage diseases, and insect defoliation. Fusiform rust continues to be the most serious disease in this biome. Approximately 6.8 million ha of forest were affected. Annosus root disease continues to be the most serious root disease in the southern regions, weakening trees and making them susceptible to pine bark beetles. Pollution Symptoms In 1990, sulfate and nitrate deposition levels were highest in the eastern United States. Ozone levels over a 10-year period were highest in the mid-Atlantic and upper southeastern coastal plain states. Black cherry and sweetgum were damaged on three quarters of monitoring plots. Over 60% of black cherry, sassafras, and yellow poplar were damaged in the upper and middle of the crowns. Ozone injury was recorded from late May in the southeast part of the biome to mid-July in the northeastern part of the biome. Ozone caused greater damage to species in the southeast part of the biome because of higher exposures for longer periods compared with northern areas. The trees have longer periods of susceptibility because they flush earlier. The higher humidity levels in this biome also facilitate gas exchange and the uptake of ozone into the leaves. In summary, this biome has a wide variety of economically and aesthetically important tree species affected by a wide variety of insects, diseases, and climatic factors. Pollution damage affects the forests regionally and causes considerable damage to the more sensitive species. 3. Mixed Mountain and Highland Systems These continental interior forests are often fire-dominated. Some harvesting, land conversion to agriculture, weather damage, and occasional pollution events also cause stress to these forests. Tree Mortality Mortality was at normal levels for most species in the northeastern part of the biome. Exceptions occurred for lodgepole pine and Douglas-fir which had higher than average mortality caused by natural thinning and root rot, respectively. 13 There is not a quantification of the mortality percentage of the Mexican forests; however, it is higher than average mortality reported for the United States and Canada and is caused by bark beetles throughout most of the country. In addition, parasitic plants, tree diseases, and insects cause significant losses. Tree Condition The crowns of most trees sampled in this biome were affected by many insects and diseases. Weather effects caused some needle damage or loss at levels considered normal for this biome. Needle flecking, a discoloration caused by snow deposition and reflected sunlight, was common at higher elevations. Needle discoloration in alpine fir was caused by various diseases, for example, Lirula abietis-concoloris (Mayr ex Dearn.) Darker, and a secondary fungus, Stegopezizella balsameae (Davis) Syd. Foliage of Douglas-fir was damaged by Swiss needle cast and snow blight. Some crown damage was recorded on lodgepole pine from atropellis canker and pine needle cast disease. Bark beetles, Dendroctonus mexicanus, D. adjuntus, and D. frontalis, caused most of the damage on mixed-coniferous forests in Mexico, where they are affecting 8644 ha. Parasitic plants such as Arceuthobium sp., Psittacanthus sp., and Phoradendron sp. are present on conifer and broadleaf trees. The first is known as dwarf misletoe and is affecting 1.8 million ha just in Mexico. These tree mistletoes are widely distributed in the biome. Other foliar injuries are caused by defoliating insects: Evita hyalinaria, Zadiprion sp., Neodiprion sp., Baliosus sp., Malacosoma incurvum azteca, and Pterophylla. There are also borer insects of branches and buds such as Stylosus sp., Eucosma sp., Rhyacionia sp., Conophthorus sp., and Paranthrene dolli. The fungus diseases affecting small areas of Mexican forests but without significant damage are Heterobasidium annosum, Phellinus pini, and Fusarium subglutinans. These diseases are considered high risk and have the potential to spread. Pollution Symptoms In biome 3, ozone stress typically occurs in the mixedconiferous forests occurring between 1500 and 2500 m in California, and to a lesser degree some mountain forests in southern Arizona up to 4000 m. Ozone is particularly injurious to ponderosa pine and Jeffrey pine, and less injurious on black oak. Loss of needle whorls, and growth reductions were the foliar injury symptoms observed in the San Bernardino and Sierra Nevada mountains. In this biome, sulfate emissions peaked in the early 1960s and by the early 1980s peak levels decreased by 20%. While sulfate emissions are generally decreasing, nitrate emissions have been steadily increasing. Nitrate emissions tend to be greatest near major cities in California, Washington, Colorado, and Arizona. No damage to forests has been reported. 14 Health of North American Forests Conifers are more affected by ozone than broadleaf species; Pinus hartwegii has been described as the most sensitive species showing chlorotic mottling, banding, severe defoliation, branch, mortality, and poor natural regeneration. Other species affected include Pinus patula, P. montezumae, P. teocote, P. radiata, and Abies religiosa. In the Valley of Mexico, Abies religiosa shows a severe branch mortality, defoliation, and a reddishbrown color on the upper leaf surface of the older leaves. Some conifers have photochemical oxidant damage near urban areas. There is white flecking on pine trees caused by insects that resemble pollution symptoms. Different species of broadleaf trees and shrubs are evident in downwind areas near urban areas and frequently have reddish-brown stippling on the upper side of the leaves resembling symptoms of oxidant damage. 4. Subtropical and Temperate Rain Forests These forests contain the largest and oldest trees in the world. Species such as Douglas-fir, Sitka spruce, and sequoia are world famous for their size and age. Fires have been the major factor altering forest ecosystems but they have occurred rarely. More recently, drought, harvesting, and exclusion of fire have combined to render the current ecosystems more vulnerable to pests, diseases, and fires. Fire prevention programs allow a large buildup of fuel, thus increasing the risk of future fires that may be more destructive than if normal ecosystem development were allowed. Major forest pests are western spruce budworm, Douglas-fir tussock moth, mountain pine beetle, Douglas-fir beetle, spruce beetle, fir engraver, western pine beetle, and pine engraver beetle. Tree Mortality Mortality has been within the normal levels in these forests and was caused by natural thinning, attack by bark beetles, and by occasional blowdown. Tree Condition Crowns of conifer species appeared healthy and had no damage. Western red cedar and western hemlock were generally healthy and had some crown thinning, chlorosis, and traces of foliage discoloration due to crown shading, suppression, and storm damage. The western spruce budworm and storms damaged Douglas-fir. Discoloration and chlorotic lower crown foliage from Swiss needle cast and snow blight fungus also occurred sporadically. Pollution Symptoms In general, there were few symptoms of pollution observed in these forests. Some occasional needle flecking occurs in localized situations but this has not been shown to have been caused by pollution. 5. Evergreen Sclerophyllous Forests The topography of this biome is mountainous, and forests are found on lower slopes and river valleys. Forest distribution is limited largely by rainfall and soil conditions. The biome contains a wide variety of species, and conifers dominate the landscape. Major conifers include western white pine, ponderosa pine, lodgepole pine, grey pine, Jeffrey pine, Coulter pine, sugar pine, singleleaf pinyon pine, incense cedar, Douglas-fir, white fir, and mountain hemlock. Major hardwoods include California black oak, canyon live oak, interior live oak, blue oak, Oregon white oak, Engelmann oak, dogwood, big leaf maple, willow, and trembling aspen. Forests in this biome have been widely affected by forest fires which are a major factor in successional patterns. Human habitation also affects forests largely because of land use changes from forest to settlement, agriculture, and grazing. Forest harvesting occurs widely. Tree Mortality Tree mortality is frequently at levels higher than expected from natural thinning in unmanaged stands. This is caused by the effects of a variety of insects and diseases. The major insect pests affecting forests include the Jeffrey pine beetle, the lodgepole needleminer, mountain pine beetle, western pine beetle, pine engravers, black pineleaf scale, and the California flatheaded borer. Diseases affecting forests include the annosus root disease, dwarf mistletoe, black stain root disease, and white pine blister rust. Abiotic damage caused by ozone damage on pines also affects many forests. A combination of multiple organisms or an interaction of site and stand conditions and insects causes most of the mortality in commercial forests of this biome. These interactions are usually between bark beetles, dwarf mistletoe, root diseases, and overstocking after disturbance. Tree Condition Crowns of both conifers and hardwoods are affected by defoliating insects. These pests are mostly species-specific and damage trees at endemic and outbreak levels. A common stress on the trees is drought and this aggravates the damage caused by other stresses. Symptoms of ozone damage on pines are common in certain areas. Pollution Symptoms In this biome, ozone is a major factor in forest health, particularly on Jeffrey pine and ponderosa pine. In the 1960s, ozone was identified as the factor causing damage to pine forests in the San Bernardino Mountains east of Los Angeles. In 1971, the first report of ozone injury to pines in the Sierra Nevada Moun- Health of North American Forests tains was published. Damage symptoms are chlorotic mottling on pine needles. Ozone injury occurs throughout most of the Sierra Nevada Mountains, and damage increases from north to south. 6. Tropical Dry Deciduous Forests or Woodlands This is a species-rich biome in western and southern Mexico that has distinct wet and dry seasons. These forests are affected by several anthropogenic stresses, deforestation, wildfire, and grazing. Tree Mortality There is not a quantitative estimate of mortality at present. These forest types are largely free of insect and disease damage and of air pollution. Tree mortality is normal for unmanaged forests in this biome and is caused by wildfires, grazing, and land use change for agriculture and livestock uses. Tree Condition The crowns of the broadleaf species are generally in good condition. There are no important conifer species in this type of forest. Occasionally insects or diseases affect some species without significant damage. Drought and poor site conditions stress most tree species but are not important factors of damage. In southern Mexico, there is a bark beetle, Dendroctonus frontalis, attacking Pinus pringlei and P. oocarpa in the States of Oaxaca and Guerrero. Pollution Symptoms Because most of these tree species are living under dry conditions, there is not significant foliar damage. There are not specific studies on this particular kind of vegetation in Mexico, although a large area of the country is within this ecosystem. More detailed studies are needed to explain the factors affecting these forests. 7. Tropical Humid Forests This biome consists of the southern and coastal Mexican forests containing many species in their diverse ecosystems. Human influence is strong; many areas are cleared for agriculture and settlement. Pollution from point sources is common. Tree Mortality There is no estimate of mortality available in this biome. Causes of tree mortality include natural and induced thinning, as well as forest clearing for agriculture and livestock. In this biome, the heavy rains can easily cause soil erosion. Several local or regional tree diseases and insects are also important. Damage resembling that caused by sulfate (SO2) on trees in the areas is probably caused by emissions from oil refineries. 15 Tree Condition The crowns of broadleaf and conifer species appear generally healthy and have little damage. Occasionally storms and wildfires cause relatively severe injury to these forests. Some insects present are Hypsipyla grandella, Xyleborus sp., and Chrysobothris sp., attacking Cedrela mexicana and Swietenia macrophylla in nurseries, plantations, and natural regeneration; Rhyacionia frustana attacks Pinus oocarpa, P. pringlei, and P. caribaea; and Hylesia frigida attacks some species of pines. Different species of rodents, ants, and locusts are affecting nurseries and plantations. A few tree pathogens attacking these forests are Puccinia spp. on pines, and true mistletoes such as Psittacanthus sp., Phoradendron sp., and Struthantus sp. on broadfleaf species. Pollution Symptoms In general, there are no symptoms of air pollution on these forests; however, near oil refineries, damage similar to SO2 occurs on some species. Leaf chlorosis or necrosis occurs intervenally. There are no specific studies dealing with this symptomatology to confirm this particular kind of damage. Research is needed in this area of air pollution effects on vegetation in the tropical humid forests. Conclusions To protect our forests and ensure their survival and sustainability for the use and pleasure of future generations, we need to be aware of the state of their health. If this is known and if they are in danger, action can be taken to preserve and protect them. Forests are essential to the continued economic and spiritual health of all North Americans. Forests are subjected to natural and anthropogenic stresses that may cause fundamental damage and changes to forest ecosystems. Monitoring the state of the health of the forests is done by assessing the mortality rates, and the condition of the crowns to determine what if anything is damaging them. The results of these monitoring activities are used to assess the health of the forests. Tree mortality in North American forests was generally in the normal range of 1–3% and is caused largely by competition and suppression within stands. The effects of insects, diseases, drought, and storms were observed frequently, and occasionally caused higher rates of mortality. Trees in some areas improved and recovered from previous damaging stresses. Defoliation and dieback of tree crowns were observed frequently and were caused primarily by known damaging agents: insects, diseases, drought, frost, etc. Classic symptoms of air pollution were found in some forest ecosystems. Damage from acid fogs and ozone occurred on several species in several 16 Health of North American Forests biomes indicating the potential for extensive damage in the absence of corrective action. In addition, trees might have been weakened or stressed by air pollution and this stress may not yet be evident in the monitoring programs. Overall, the assessments indicate that there is no large-scale decline in the health of forests that can be attributed solely to atmospheric pollution. Nevertheless, continued monitoring is necessary to ensure that the health of these forests is protected. This publication is a general overview of the situation; for more detailed results, the reader is referred to the national publications on the subject. Bauer, L.I., de; Hernández-T., T.; Manning, W.J. 1985. Ozone causes needle injury and tree decline in Pinus hartwegii at high altitudes in the mountains around Mexico City. J. Air Pollut. Control Assoc. 8: 838. Bauer, L. I. de; Krupa, S.V. 1990. The Valley of Mexico: Summary of observational studies on its air quality and effects on vegetation. Environ. Pollut. 65: 109–118. Becker, M.G.; Landmann, G.; Levy, G. 1989. Silver fir decline in the Vosges mountains [France]: role of climate and silviculture. Water Air Soil Pollut. 48: 77–86. 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