Wildlife Considerations in Modern Forest Management
By John P. Hayes
Professor of Wildlife Ecology, Oregon State University

In recent years, concerns over wildlife conservation have shaped forest policy and forest management decisions in the United States in ways that were almost unimaginable two or three decades ago. Species such as the northern spotted owl and marbled murrelet in the Pacific Northwest, the red-cockaded woodpecker in the Southeast, and the Indiana bat in the Northeast and Midwest were once largely known only to wildlife enthusiasts, but now drive policy and management decisions in many forests because of their listing as threatened or endangered under the federal Endangered Species Act.

Although the current attention to wildlife and emphasis on biodiversity in forests has resulted in abrupt changes in forest management practices, the seeds of change that led to our current approach to managing wildlife in forests were planted quite some time ago, and close interactions between the fields of wildlife and forestry have a rich history. Until relatively recently much of the attention that forest managers gave to wildlife in forests focused on managing habitat for game species and on minimizing impacts of wildlife on seed production and forest regeneration.

The 1960s and 1970s saw the beginning of a transformation in our perspectives on wildlife-forestry interactions, coinciding with increased public awareness of environmental issues and passage of key legislation such as the Wilderness Act, the National Environmental Policy Act and the Endangered Species Act. Concurrent with this new view was a proliferation of studies on the ecology of wildlife in forests and enhanced understanding of the importance of dead wood, riparian habitat, old-growth forests and complex forest structure for wildlife.

A central paradigm emerging from this research relates to the concept of spatial scale. A large number of recent studies have demonstrated that wildlife select different habitat characteristics at different spatial scales, and that attention to forest characteristics at different scales is critical to provide high quality habitat for wildlife. For example, early attempts to provide habitat for northern spotted owls emphasized protection of the nest tree (habitat component scale). We now understand that the characteristics of the stand in which the nest tree occurs are also important (stand scale), as is the amount, distribution and characteristics of stands surrounding the nest stand (landscape scale) and connectivity among different nesting areas (regional scale). To effectively manage for this species, attention to each of these scales is necessary.

Within forest stands, key aspects of habitat to which wildlife respond include vertical and horizontal heterogeneity, amount of standing and fallen dead wood, and vegetative species diversity. Special habitat features, such as riparian areas, rocky outcrops and unique plant associations, also provide important habitat for a number of species of wildlife. Forest management that maintains or increases structural complexity within stands can provide multiple niches for wildlife.

For many species of wildlife, standing dead wood, or snags, is particularly important. Woodpeckers and a suite of cavity-nesting birds, bats, forest carnivores, arboreal rodents, and other species are dependent on or highly associated with snags. Few management decisions influence abundance and composition of wildlife communities as strongly as decisions concerning management of snags. Maintaining large diameter snags in a variety of stages of decay is a key management strategy to providing habitat for several species.

Just as many species have been shown to prefer the complex forest structure found in old-growth stands, others are closely associated with stands in early stages of development. Because of the diversity of habitat needs of different wildlife species, maintaining a wide array of stand conditions helps promote a diverse wildlife community. A good example of an approach to maintain a variety of structural conditions through time is the recent management plan developed by the Oregon Department of Forestry. This plan manages for different structural conditions spread across the landscape, providing connectivity between key habitat types. Appropriately, these plans call for maintaining a minimum amount of habitat in the stem-exclusion stage, as this developmental stage tends to be poor quality habitat for most wildlife species.

While providing the diversity of habitat to meet the needs of wildlife species can be a daunting task, even simple steps, such as providing dead wood in forest stands, managing for a variety of stand conditions, and protection of special habitat features, can result in significant benefits for wildlife.

Extracted from: Society of American Foresters, Western Forester 49(4) 2004


Wood Decay in Healthy Forests: The Paradox and the Promise
by Bruce G. Marcot
USDA Forest Service, Pacific Northwest Research Station

A Legacy of Snags in the System
It is widely acknowledged that standing dead or partially dead trees provide important habitat for woodpeckers and other primary cavity excavators, and that a wide array of other animals use woodpecker cavities. Guidelines exist for maintaining or even creating snags on federal lands, state forest lands and private forest lands. It is also known that large-diameter snags held over from previous, older stands constitute one aspect of “wood legacies” that provide ecological functions more fully than small-diameter snags produced from understory suppression during even-age stand management.

Snags are utilized by wildlife in many ways. For example, red squirrels use witches’ brooms created by mistletoe in Douglas-fir trees. Black bears den in hollow trees and logs in northeastern Oregon. Shredded down wood and bark piles at the base of snags are often used by salamanders in Douglas-fir forests of Washington. Long-eared bats use tree stumps for roosts. High-cut stumps are used by white-headed woodpeckers for cavities. Ponderosa pine snags are used as breeding roosts by bats.

However, it has only been more recently acknowledged that many other forms of wood decay—other “wood decay elements” or WDEs—than just snags also provide vital ecological services in forest, woodland and riparian ecosystems. This is a brief review of the diversity and ecological roles of WDEs.

Beyond snags: elements and functions of wood decay elements
WDEs also include down wood, root wads, tree stumps, litter, duff, broomed or diseased branches, hollow trees and partially dead trees. They also provide resources and substrates for many organisms that perform vital ecological roles of transforming and cycling nutrients, decomposition, respiration and other biological processes. Such roles benefit ecosystems far beyond the confines of the wood decay elements per se, and are a natural and vital part of native forests and ecosystem processes. Providing for WDEs may seem like sacrificing growing space, but in the long run, ecological processes of WDEs greatly contribute to overall ecosystem health, soil productivity and growth of desired tree species.

Beyond just providing wildlife habitat, WDEs also provide for an array of ecological roles of wildlife in forest ecosystems. For example, in southwest Oregon conifer-hardwood forests, hollow trees are used by 24 wildlife species, including nine bats, five owls, two woodpeckers, a swift and others. Of these 24 species, five (two birds and three mammals) also serve the beneficial function of dispersing seeds and fruits of native plants, and two (mammals) tunnel in soil that in turn creates burrows used by other species and can help improve soil structure and uptake of organic matter. These and other secondary beneficial ecosystem services are provided by many wildlife
species associated with all WDE categories in all forests.

Down wood has a high pore volume and serves as moisture reservoirs, and in moderation, provides microsites for beneficial mychorrizal fungi, plants and animals that aid in forest recovery after prolonged drought or fire. Large pieces of down wood eventually work into the soil where they serve as long-term, time-release sources of humus, organic matter, phosphate and nitrogen, and mediate soil nutrient cycles. Woody material in the soil creates acidic soil conditions that favor soil microbial activity that help fix nitrogen for use by trees. Down wood on the surface can help stabilize soil movement and deter erosion, and serve as nurse logs for spruce, hemlock, alder, Douglas-fir and other trees. Wood that enters the soil profile tends to stay there. Some carbon dating studies in the Inland West have shown buried wood material to be 500 to 1,000+ years old. All this means that restoring natural levels of coarse wood in soil horizons may be an immensely long-term process.

The paradox of health and harm
Such direct and indirect benefits of WDEs, however, are often ignored because of the fear that WDEs (especially snags and down wood) increase undesirable fire and insect hazards. In some situations this is true, but the ecological benefits of WDEs contributing to healthy, diverse, productive forests over time could be weighed against any such short-term risks. The challenge to forest managers is to provide enough
WDEs in a variety of types and spatial patterns for long-term soil productivity, ecosystem functions, and use and need by organisms, but at the same time to avoid undue hazards of fire, insect pest outbreaks, and operational and recreational safety problems.

The promise of how much, where and how
WDEs could be provided in both clumped and dispersed patterns, particularly taking advantage of natural patterns such as small, isolated root rot pockets that create local clumps of snags and hollow trees, or low-quality trees with splitting or sloughing bark growing on rocky soil or sites unfit for commercial tree use. More widely dispersing rather than clumping WDEs in fire hazard zones, such as urban-forest interfaces, may provide some measure of wood decay, but not contribute to fire and insect pest risks.

Also, managing for WDEs can be evaluated at stand, local watershed and broader landscape scales to ensure that large areas are not devoid of legacy wood elements. Most important, by considering WDEs in forest management, managers are planning for the future when they account for dynamics of tree growth and decay, stand rotation and disturbance dynamics (particularly fire, insect pests and pathogens).

How much WDEs should be provided? That depends, of course, on the overall forest management objectives. One possible objective is to provide a minimal amount of WDEs in more intensive tree farms to help complement greater amounts of WDEs found on adjacent national forests or other multiple-use forest lands. Other possible objectives may include providing greater patches of WDEs on less productive growing sites, or rotating stand management methods across the landscape so that each site gets an occasional infusion of WDE legacies and large wood into the soil. To various degrees, the forest manager may wish to emulate some range of historic or unharvested conditions.

In forests of the Inland West, about 30 percent of organic volume of soils will maintain peak mycorrhizae amounts in the organic soil horizon. This translates to about 10-15 tons/acre of surface down wood, which should be relatively large woody residue scattered across areas with minimal soil disturbance. Chipping of fuel wood is not an ecologically viable solution, as rainfall leaches large amounts of toxic, water-soluble phenolics from the chips and blocks soil structure, causing mortality of tree seedlings. In some cases, artificial logs could be introduced to cover greater than 25 percent of the area in piles large enough to provide deposits of large coarse wood similar to natural levels.

Broadening the vision
The overall lesson is that the forest ecosystem manager can view WDEs as natural and desirable parts of the forest ecosystem, and more than just providing occasional snags for woodpeckers. All types of WDEs provide important functions for plants, animals and ecological processes that together maintain natural, diverse and self-sustaining forests.

Extracted from: Society of American Foresters, Western Forester 49(4) 2004


Vertebrate Use of Dead Wood in the Pacific Northwest
by Mark Boyland & Fred L. Bunnell (2002)
University of British Columbia Vancouver, Canada


Abstract: The many unique features of dead wood make it attractive as a place to forage, nest, and shelter. In the Pacific northwest, 69 vertebrate species commonly use cavities, and 47 species respond positively to down wood. Cavity users typically represent 25% to 30% of the terrestrial vertebrate fauna in forests of the Pacific Northwest. Cavity excavators select for heart rot—major factors governing it’s presence are tree species, site productivity, tree age, and time. Data presented shows 2-3 large snags per hectare, and 10-20 smaller snags per hectare, throughout the rotation, are required to sustain cavity nesting birds.


Ecosystem Components that Encourage Wildlife Diversity
Extracted from
Appendix: Wildlife Habitat Relationships, Coast Range Forests
Publication R6-NR-ECOL-TP-03-02.
USFS Pacific Northwest Region, 200

Dead and partly decomposing trees
     Approximately one-third of bird and mammal species in forested landscapes use tree cavities for denning, nesting, or roosting. Snags, dead tree tops, and otherwise decayed portions of live trees provide opportunity for woodpeckers and other species to create holes. These cavities are in turn used by secondary cavity-nesters that search for and use these cavities, rather than create their own. Cracks, crevices and loose bark also provide nesting and roosting substrates for bats and brown creepers. Probably the rarest structures in the forest important to vertebrates, and the most difficult to duplicate, are large hollow trees. These are often western redcedar or incense cedar, but can be just about any species of tree. Some trees are hollow from the bottom up, some from the top down, some only in the middle (but the latter are very difficult to find). Bears, bats, swifts, and other mammals and birds utilize these structures. Vaux’s swifts nesting in forests exclusively use these structures.

Down wood
     Logs are used by a wide variety of wildlife, but small mammals and amphibians are probably the groups most dependent upon these structures. Some species prefer more sound structures, utilizing the space created by loose bark, while others predominate in more decayed structures that are soft enough to tunnel through or that have a matrix of navigable cracks due to the work of brown cubical rot. Many species utilize logs simply for hiding cover, nesting cover, travelways, or perches.

Diversity of tree species
     Different tree species germinate in different ground conditions, grow at different rates, exhibit different shapes to their crowns, boles, and leaves, have different susceptibilities to root rots, stem rots, and mistletoes, are differentially resistant to stem breakage from wind, ice, and snow, attract or repel different communities of invertebrates, and finally, different tree species have different maximum heights and senesce and die at different ages. All these differences suggest that a wide variety of nesting, foraging, roosting, hiding, and resting habitats may be produced by different combinations of species, ages, and conditions of trees. Thus, single-species stands typically exhibit less vertebrate diversity than multi-species stands.

Broadleaf trees
     The significant differences between conifers and broadleaf trees cause the combination of these species in forest stands and landscapes to significantly increase the number of animal species present. Warbling vireos are most frequent in areas with abundant broadleaf trees or tall shrubs, and black-throated gray warblers and black-headed grosbeaks prefer mixed habitats. In the Douglas-fir and drier western hemlock associations on the eastern foothills, broadleaf trees (especially oak) attract western gray squirrels, but in all plant associations, sites dominated by broadleaf trees are likely absent of Douglas squirrels. Broadleaf trees may be important to mollusk diversity.

Shrubs
     Forest understory shrubs provide nesting structures for Swainson’s thrushes, winter wrens, and Wilson’s warblers. The Wilson’s warbler in particular utilizes tall deciduous shrubs for nesting. Shrubs provide important habitat for invertebrates, browse for deer and elk, and cover for a wide variety of birds, mammals, and reptiles. Patches of older shrubs in particular can be hotspots for arthropod, lichen, and bryophyte diversity.

Fruits, berries, and nuts
     Numerous trees, shrubs, and forbs produce seeds and soft fruits that are consumed by a wide variety of birds and mammals. Some of the more common species producing mast include all conifers, maples, and hazel. Oregon white oak and Pacific madrone occur sparingly, most often in or near the Douglas-fir series. Species producing berries (i.e. seeds with a fleshy outer layer) include dwarf and tall Oregon grapes, salal, several species of blackberry, thimbleberry, several species of manzanita, salmonberry, bitter cherry, snowberry, several species of rose, Pacific dogwood, several species of huckleberry, blue and red elderberry, and cascara.

Soil and forest litter
     Soil characteristics combine with annual temperature, precipitation, and solar exposure to determine suitability and growth potential of a site for plant species and communities. Burrowing animals such as gophers, moles, some voles, and mountain beaver prefer relatively porous soil. Several species of shrews appear to be particularly abundant where forest floor litter is abundant and deep, and western red-backed voles are especially common in areas with a thick duff layer. Forest floor characteristics are also important to ground-dwelling invertebrate communities.

Rocks, cliffs, caves
     Many different structures are created by rock. Cobble-sized talus is common below cliffs and on steep rocky slopes. These habitats may be dominated by several species of amphibians (e.g. western redback salamander, clouded salamander) if wet, and several species of snakes (e.g. northwestern garter snake) and lizards (e.g. northern alligator lizard) if dry, and some communities have both. Mice and voles also inhabit talus. Accumulations of larger rocks provide homes for long-tailed weasels and potential denning sites for other medium to large mammals. Cliffs provide nest sites for peregrine falcons, common ravens, and violet-green swallows, and caves are often home to turkey vultures, bats, and medium and large mammals. In arid associations, seepy areas provide rare habitats for amphibians and mollusks.

Water
     While many animals gain a substantial portion of their water needs from the food they eat, most also require consumption of water on a near-daily basis. While small mammals and birds can often obtain water from condensation on vegetation, larger animals are more dependent upon more substantial water sources such as streams and ponds. These undoubtedly provide the predominant water source for the majority of birds and mammals in most landscapes. Even migratory birds, presumably unfamiliar with any particular small stream, have an uncanny ability to locate small trickles and pools located on an otherwise dry stream segment under a tall forest canopy.

Streams
     Pacific and Cope’s giant salamanders, Columbia and southern torrent salamanders, and tailed frogs are restricted to breeding in cool, running water. The most common of these, the Pacific Giant Salamander, contributes significantly to the biomass and predation within small streams. While terrestrial densities of stream-breeding amphibians decrease with distance from streams, and relatively high rainfall and forest floor humidity gives more freedom for terrestrial travel in western Coast Range forests.

Ponds
     Western toad, Pacific tree frog, red-legged frog, bull frog, northwestern salamander, long-toed salamander, and rough-skinned newt are restricted to breeding in still or very slow-moving water. Highest terrestrial densities of these species are found in close proximity to breeding sites, and sometimes concentrate on particular hillsides or along particular inflowing or outflowing streams. Some species exhibit substantial dispersal capabilities and may be found in very small numbers several miles from any suitable breeding habitat. While most species typically breed in ponds or lakes, others (e.g. Pacific tree frog) will sometimes breed in very small water, even puddles in abandoned roads, and roadside ditches in open roads. Some require semi-permanent water; for example, northwestern salamanders require more than one season to metamorphose, and may even become neotenic (a permanent “larval” form that is capable of reproduction), while others (e.g. Pacific tree frog) can breed in water that dries up during summer, because they metamorphose rapidly. Some (e.g. northwestern salamander, red-legged frog) require substrate such as sedges or woody plant stems for egg-laying, while others (e.g. Pacific tree frog, rough-skinned newt) do not require such substrate, though the newt may have requirements for pond-bottom composition.
     The marsh shrew and wood duck are common inhabitants of ponds as well as small streamside wetlands. Common garter snakes in particular are attracted to ponds containing frog tadpoles. Marsh shrews also take advantage of this abundant food source. Vaux’s swifts, common nighthawks, and several species of swallows and bats obtain water in flight at ponds and still water pools in streams and rivers. These species also forage on flying insects over these waters.