Ecology of Ungulates: A Handbook of Species in Eastern Europe and Northern and Central Asia

A Handbook of Species in Eastern Europe and Northern and Central Asia The knowledge on the ecology of ungulates (orders Perissodactyla and Artiodactyla).
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The now recovering temperate forests constitute a large and growing source of carbon storage, wildlife habitat and watershed protection. Many important products come from the temperate forest, including sources of non-timber forest products wild plants and fungi used as food, hunting animals for food, maple syrup [N. America] from Acer saccharum, and birch sap for traditional beverages and in folk medicine as an ingredient for different antiseptic and anti-inflammatory treatments [Central and NE Europe] , wood products hardwood used for furniture and flooring, Pinus and Quercus used for construction lumber , and paper products many species used for pulp.

The forests also contribute greatly to the tourism industry through attractions such as remnant old-growth forests, large trees, spring wildflowers, brilliant fall colors, and habitat for migrant bird species, song birds, and other wildlife species commonly viewed by tourists. Soil structure in temperate forests usually has an organic horizon at the top of the soil profile composed of leaf litter in various stages of decay, from fresh leaves at the top to fragmented leaves in the middle and black humus at the bottom. Rate of decay is important for determining how fast nutrients are released from the organic horizon, and is determined by temperature and soil moisture, which in turn is determined by climate and soil texture.

Organic horizon thickness is greater in colder climates and drier climates within the temperate zone, where decomposition of leaf litter is slower, in forests dominated by species with high C: N ratios and low nitrogen content of leaf litter Fagus , Picea , Pinus , Quercus , in forests on sandy soils, and forests with low earthworm abundance.

Cold temperatures in northern temperate and boreal forests depress organic matter decomposition, and may suppress nutrient movements in soil and their uptake by roots. Young, postglacial soils tend to be more N- than P-limited Reich and Oleksyn Temperate old-growth forest ecosystems are particularly vulnerable to increases in atmospheric N deposition.

In response to increased N supply both from N deposition and faster litter mineralization enhanced by increasing temperatures , the pattern of N cycling may change and be accompanied by increased selection favoring nitrophilous plant species and a decrease in ground species richness Tamm An example of large-scale changes in nutrient cycling and soil function caused by human activity has occurred in Europe during the late nineteenth and twentieth centuries.

Coniferous trees including native Picea abies, Pinus sylvestris , and North American Picea sitchensis , Pseudotsuga menziesii species were widely planted to reverse eighteenth and nineteenth century deforestation and increase productivity, often replacing native deciduous species. Major disturbances that kill or level the forest canopy include logging, wind, fire, ice storm, landslides, and insect infestation.

Note that all of these may also create partial disturbance with scattered tree mortality contributing to the gap dynamics of multi-aged stands described below. Whether stand-leveling disturbance initiates an episode of succession—defined as a directional change in species composition over time, usually establishment of a dominant cohort of early-successional pioneer trees species followed by its gradual replacement by a suite of later-successional species—is context dependent Heinselman , Finegan For example, canopy-leveling wind storms or logging of the large trees may leave a carpet of late-successional advanced regeneration intact on the forest floor, leading to initiation of an even-aged stand followed by a stand development sequence from even-aged to multi-aged, but not necessarily change in species composition.

This has been shown to occur in North American Acer saccharum and Tsuga canadensis forests. On the other hand, high-severity disturbances such as windthrow followed by high-intensity fire, or landslide could easily wipe out the advanced regeneration, leading to establishment of shade-intolerant, early-successional species succeeding to shade-tolerant, late-successional species over the subsequent one to two centuries.

Illustrative case studies include Betula papyrifera and Populus tremuloides succeeding to Acer saccharum and Tsuga canadensis in North America and Betula grossa succeeding to Tsuga sieboldii and Fagus crenata in Japan Yoshida and Ohsawa , Frelich Multiple disturbance regimes occur in the northern temperate forest biome. The least severe regime occurs on good quality sites with moist climates, and is dominated by single to multiple treefalls, usually wind-dominated regimes with some insect, disease, and drought mortality, where gap dynamics are prominent described in more detail below.

Such forests are dominated by mid-to-late-successional species, and under natural conditions, episodes of succession depend on rare episodes of severe disturbance mentioned above. However, in the last one to two centuries, clear cutting followed by slash burning has mimicked the rare natural wind—fire combination and created much larger expanses of early-successional forests than would have occurred naturally, in regions undergoing human settlement. These forests are also multi-aged and have little succession over time unless the natural disturbance regime is disrupted by humans, usually by fire exclusion.

Once established, these trees commonly live a century or more and survive many subsequent fires.

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These forests occupy relatively dry climates in the transition from temperate forests to grasslands or Mediterranean vegetation, as well as excessively well drained sites on sand plains and south-facing rocky hillsides within wetter climates. A third disturbance regime is that of mixed-intensity fire, whereby fires of variable intensity occur at irregular intervals.

Fires may kill some, most or all trees in a given stand, and may come at short intervals so that there is no time for succession, maintaining the current successional status, or long intervals, allowing succession to take place between fires. Although successional pathways for these forests are understood, i. Large tracts of temperate forest occur on mesic sites where stand-leveling disturbances are rare, and gaps from single to several trees falling constitute the main type of disturbance e.

The dynamics of these treefall gaps, canopy turnover rates, canopy tree residence times, canopy structure, and recruitment of new trees into gaps have been studied at numerous sites. Such forests are usually multi-aged, and dominated by late-successional, shade-tolerant species, with some mid-tolerant species, and an occasional specimen of early-successional species. Fluctuations in relative abundance of the late-successional species occur over time due to influences on seedling success such as preferential deer grazing on seedlings, and neighborhood effects structure and chemical properties of the leaf litter, sprouting; Brisson et al.

The late-successional species generally exist as suppressed seedlings or advanced regeneration that will record a release from suppression if the tree above dies; this fact has been used to reconstruct the treefall history of forests going back as much as several centuries Lorimer , Payette et al. In contrast, the seedling bank of mid-tolerant tree species is rather small due to rapid turnover as seedlings die due to low light levels, but these species are adapted to enter new gaps by seed dispersal followed by rapid height growth that may surpass that of shade-tolerant species.

These gaps can have not only more light, but higher soil and air temperatures, and more water due to lack of use by the fallen tree , leading to more nutrient availability in the first few years as the fine roots of the fallen tree decay in the relatively warm, wet environment. Tree species mid-tolerant of shade e. Fraxinus and Tilia , along with some species of Acer , Betula , Carya , Pinus , and Quercus often take advantage of these gaps, which therefore help to maintain local species richness over time.

In addition, treefalls create coarse woody debris, which, when it reaches an advanced state of decay, can also be a microsite for germination of some tree species that cannot tolerate thick leaf litter or the dense herbaceous vegetation that may cover the forest floor. These decaying logs often last 20—40 years for angiosperm species in genera such as Acer, Quercus and Tilia , but may last 1—2 centuries for gymnosperms such as Picea, Pinus and Tsuga.

Decaying logs also are small-scale hot spots of biodiversity due to the large number of species of mosses, fungi, insects, and amphibians that use them as habitat due to their relatively high water content Harmon et al. Interactions of note in the northern temperate forest include the occurrence of mosaics formed by patches of evergreen conifer and broadleaf deciduous species, shade-tolerant and intolerant species interactions, and the presence of numerous understory species that take advantage of a brief period of combined warmth and light in spring, and heterotrophic species that do not require sunlight to fuel carbon gain e.

Evergreen—deciduous mosaics of Tsuga or Picea with Acer , Betula and Tilia mesic sites , or with Pinus and Quercus drier sites occur in the parts of the temperate forests with cooler climates, including the northern USA, northern Europe, and Asia. These mosaics Figure 3. Shade tolerance or avoiding shade is necessary for any plant species to reproduce in these forests given dense canopies and forest floor light levels ranging from 1—10 percent of full sunlight.

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Plant species have evolved a variety of strategies. Spring ephemerals are species that grow rapidly as soon as winter ends, taking advantage of the sunlight available on the forest floor prior to canopy leaf out. Some of these genera have vicarious species related species that live in similar environments with similar ecological niche among the major occurrences of temperate forest, for example, Erythronium americanum in the eastern USA, E. Other early blooming species are commonly misclassified as spring ephemerals, but they keep their leaves for much of the summer—these include geophytes such as Trillium , and herbs such as Arisaema jack-in-the-pulpit , Sanguinaria bloodroot , and Viola violets , which can use energy stored in a bulb or rhizomes to produce leaves early on in full sun, but continue to photosynthesize in deep shade after canopy leaf out.

Saprophytic plants such as Corallorhiza coral root orchids , Monotropa uniflora Indian pipe and parasitic plants such as Epifagus virginiana beech drops and Conopholis americana squaw root get their energy from decomposing organic matter or attachments to tree roots, and do not require sunlight. A number of plant species take advantage of treefall gaps, where higher than average light levels last a few decades, to grow up into the canopy e.

Betula , Quercus or to complete their life cycle and create long-lived buried seeds that await the next gap a century or more later—e. Prunus pensylvanica pin cherry , Rubus raspberry , and Geranium bicknellii. Epilobium angustifolium , fireweed. Shade-tolerant shrubs in forest understories can create complex spatial dynamics for tree regeneration. Understory dwarf bamboo Sasa spp. For example, beech Fagus crenata and maple Acer mono seedlings in Japanese forests are restricted to small patches where bamboo is absent Yamamoto et al.

Many symbiotic and antagonistic plant—animal interactions occur in northern temperate forests, involving seed dispersal, pollination, defoliation, insect herbivory, and preferential grazing by large mammals deer, moose that can alter successional trajectories. Although many species of temperate trees have wind dispersed seeds, animal dispersal zoochory also occurs.

One well-known case is for acorns of northern red oak Quercus rubra that are cached by various animal species in the fall for later use during winter. Gray squirrels Sciurus carolinensis bury acorns in the soil and are known to remember the locations of several thousand acorns, but during mast years, they often bury more than they retrieve, leaving many to germinate, and at the same time hiding those acorns from consumption by deer Odocoileus virginianus , black bear Ursus americanus , and turkey Meleagris gallopavo.

Blue jays Cyanocitta cristata can carry acorns, and commonly fly several hundred meters to two kilometers from the parent tree, scatter hoarding thousands of acorns across the landscape Johnson and Webb In Europe, acorns of Quercus coccifera , Q. Such estimates provide a good example of the value of management strategies securing critical breeding and foraging habitats of seed dispersal animals.

Despite the fact that the Eurasian jay prefers acorns of native Quercus robur it is also efficient in dispersing invasive North American Q. Increase of red oak leads to significant reduction of native species abundance and diversity Woziwoda et al. Seed dispersal by ants is known as myrmecochory, and is common among temperate forest understory plant species Gomez and Espadaler The spring wildflower genera Trillium North America and Asia and Viola North America, Europe and Asia are examples, with a number of species dispersed by ants, although other vectors also exist.

The seeds of these species have a fat-rich elaiosome attached to each seed, which the ant can detach for consumption, commonly after moving the seed one or two meters. A large number of bee species occur in temperate forests often 50 or more species in one stand , which pollinate most of the forest understory species with brightly colored flowers and limited pollen production, and a few of the tree genera such as Acer in part, wind dispersal also occurs , Prunus , and Tilia.

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Folivorous insects are common, with many species that cause ongoing low-to-moderate levels of defoliation, that interact with plant defense compounds in leaves such as phenols and terpenoids. Other insect species have periodic outbreaks every decade or longer, during which the forest may be almost totally defoliated.


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For example, deciduous forests in eastern North America can be defoliated by saddled prominent moth Heterocampa guttivitta caterpillars during droughts that prevent trees from defending themselves against insects, and on a more regular 10—12 year cycle by the forest tent caterpillar Malacosoma americanum Horsley et al. These defoliation events usually cause scattered tree mortality, especially to older trees with other health problems, but seldom cause extensive tree death at the landscape scale.

Much more serious are growth losses and tree mortality after outbreaks of the common pine sawfly Diprion pini. Defoliation by this late season feeder causes higher losses than herbivory by early season feeders, e. During severe outbreaks in Finland, approximately , ha were defoliated by D.

Grazing animals, especially ungulates like deer, elk and moose can regulate plant community composition and direct succession through their plant species preferences. Many ungulates in the temperate zone consume woody plants during winter and herbaceous plants during summer. If the deer to plant ratio is high, they can regulate the balance between relatively palatable and unpalatable plant species.

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The strength of these influences on composition is in turn embedded in a trophic cascade. For example, wolves in Wisconsin, USA create a patchy distribution in deer density across the landscape, due to predation and deer avoidance of wolf pack territories. This in turn influences lushness, composition, and species richness of herbaceous plants Callan et al. Soil animals such as earthworms, beetles, and many other taxonomic groups whether native or invasive , also influence plant community composition by altering the structure of the organic horizon, moving seeds to different layers within the soils, consuming seeds, and regulating water and nutrient cycles within the soils.

There are considerable ecological cascades within the soil Eisenhauer et al.

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The temperate zone has historically had high human populations, especially in Europe and Asia, and for the last century in North America as well. Therefore, much of the forest has been converted to croplands and cities, and almost all that remains forested has been logged at least once. In Europe and Asia most forests have been logged multiple times and planted regeneration is used on a widespread basis to supplement natural regeneration after harvesting, while in North America most forests have been logged once or twice, and natural regeneration is much more common.

In recent decades, the wildland—urban interface, an area with widely scattered isolated houses, has encroached on large tracts of forested land, creating numerous foci for introduction of invasive species, and fragmenting the landscape, favoring certain species that do well along forest edges. A very small amount of primary temperate forest remains as compared to tropical and boreal forest biomes, commonly less than 1 percent of the original forest Frelich Most of these primary remnants have been identified by conservationists and set aside in well-known preserves.

These primary forest remnants serve as templates for restoration of secondary forests, and as a baseline for the occurrence of ecosystem processes as compared to forests that are harvested. However, in many regions with thousands of years of human influence, no natural templates for restoration exist. In such cases a multi-disciplinary synthesis of historic records, silvicultural and paleoecological evidence may be needed to develop management techniques that mimic disturbances and other conditions needed to maintain the diversity of native tree species and smaller species dependent on them, and maintenance of certain cultural features of the landscape may also become a priority Lindbladh et al.

Fragmentation manifested as small woodlands of a few to a hundred hectares surrounded by agricultural lands or cities, is common in the temperate forest biome throughout the eastern USA, Europe and Asia Wilcove et al. This has led to conservation problems including: Biological Reviews 80, Breeding endangered cervids in captivity.


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