Ecosystem Diversity 595 TF ‘Species richness 8 34 5 6 Canopy dominance class Figure 20.8. Relationship between overstory and understory canopy coverage and ground-cover species richness (number of species) in moraine landforms of the University of Michigan Biological Station, northem Lower Michit- gan. Points represent plots taken in nine ecosystems in the Interlobate and Colonial Point moraines. (After Pearsall, 1995,) Beta diversity of ground-cover species, based on paired comparisons of ecosystems in upland landforms, is essentially the same for ground-cover vegetation of ecosystem types of outwash plains and moraines. Beta diversity is relatively low in the Interlobate and Colonial Point moraines where there are relatively few ecosystem types and they have similar species richness and coverage. Overall, both alpha and beta diversity analyses and relationships reveal the useful- ness of hierarchical classification and mapping of landscape ecosystems in examining biodiversity at multiple spatial levels. Although much of the outwash plain landscape ap pears relatively flat and strongly dominated by aspens, the multfactor-multscale land scape ecosystem approach reveals site differences across the landscape that are ecologi- cally meaningful and that ate reflected in differences in ground-cover vegetation, These differences are also reflected in ecosystem processes such as nutrient cycling (Zak et al. 1986) and succession (Host et al, 1987). Ecosystem Diversity In the previous examples, we have considered species diversity. In landscapes where whole ecosystems of a given scale have been distinguished and mapped, the map itself (see Figures 2.12 and 13.11) provides a visual estimate of ecosystem type richness and evenness. For example, in the Huron Mountains in Marquette County of Upper Michigan, the Pine River flows through a large expanse of Lake Superior beach (Figure 20.9). In this landscape, six riverine ecosystem types are tightly clustered along the river, and their ‘number, size, and spatial distribution are in marked contrast to the single, fat, fire-prone beach ecosystem dominated by jack pine. The riverine ecosystems exhibit diverse soil conditions that range from extremely acid and infertile peat to very fertile muck and a re- ‘markable richness of tree, shrub, herb, and bryophyte species (Simpson et al, 1990). Sim-596 Chapter 20 / Diversity Figure 20.9, Landscape ecosystem diversity. Comparison of the number, size, and pat- tem of landscape ecosystem types of an area along the south shore of Lake Superior, Huron Mountain Club, Marquette Co., upper Michigan. In con- trast to the extensive beach terrace (ecosystem type 2) six small ecosystem types (13, 18, 41, 43, 44, 50) border the Pine River where it flows through the beach ecosystem. (After Simpson et al., 1990.) ilar “hotspots” of high ecosystem diversity were easily identified on the ecosystem map of the 4,000-hectare Biological Station tract described above (Pearsall, 1995). High biodiver- sity is associated with these hotspots of ecosystem diversity in the glacial terrain of Michi ‘gan (Simpson et al., 1990; Pearsall, 1995). Where ecosystem maps are available, land ‘managers have the opportunity to efficiently conserve biodiversity by identifying areas of high ecosystem diversity. ‘The diversity of landscape ecosystems and landforms can be assessed quantitatively using conventional alpha and beta diversity indices. Also, ecosystem diversity can be quantitatively determined by using the information dimension (d.) of Lochle and Wein (1994). Using this approach, Pearsall (1995) used the number of ecosystems, multiple characteristics of their physiography, soil, and vegetation, and their pattem in the land- scape to determine the ecosystem diversity of landforms and ecosystems of the Biological Station tract. Diversity of ecosystems was highest in outwash plains, and the centers of highest diversity that were apparent from ecosystem maps were confirmed by this multi- variate diversity measure. CAUSES OF SPECIES DIVERSITY AT MULTIPLE SCALES In the following sections, we examine biological diversity from the broad scale of macro: ‘ecosystems and continental events to the local scale of microecosystems. The objective isDiversity at Continental and Subcontinental Scales 597 to consider some of the main factors that work together in creating the pattem of diversity that we observe. Diversity at Continental and Subcontinental Scales PALEOGEOGRAPHY AND CONTINENTAL RELATIONSHIPS At the macroecosystem scale, the age of continents, their relative isolation or contiguity with other land masses, and their movements across the Earth's surface have provided op- Portunity for speciation and extinction. Continental drift has had a major influence on the range of species present on any continent. Collision of continents permitted species from ‘one continent to colonize another. About 100 million years ago, the modern continents were in place, and major floras and faunas developed in a state of increasing isolation. Re- markable floras and faunas developed in Australia and New Zealand in contrast to Asia South American, Aftican, and North American floras differentiated in unique ways, but in all cases were strongly influenced by their historical origins and development. A fascinating example is the history of the beeches of the northern and southern hemispheres that share the family Fagaceae but are in different genera, Beeches of the ‘genus Fagus (“true” or northern beeches) are deciduous and differ in many other respects from the southern or “false” beeches of the genus Nothofagus. Today, southern beech forests (genus Nothofagus) are found in South America, Australia, New Zealand, New Caledonia, and New Guinea (Figure 20.10). The plant fossil record of these countries and of Antarctica shows that they might have descended from similar forest that was once ‘thought to have been extensive in the cool and temperature latitudes of the gigantic land ‘mass, Gondwanaland, that existed 150 to 100 million years ago (Poole, 1987; Barlow and ZZEXTANT Fagus 4 EXTANT Nothofague + FOSSIL Notiofagus ro sg aus anal Figure 20.10. Occurrence of the living and fossil taxa of the northern beeches (Fagus) and the southem beeches (Nothofagus). (After Poole, 1987.)9B Chapter 20 / Diversity Hyland, 1988). What the origin of those forests was and how they came to be separated from the northern hemisphere Fagus, so-called true beech—presumably true because they ‘were first named by botanists in that part of the world—remain mysteries. Figure 20.10 il~ lustrates the continental and hemispheric separation of living and fossil beeches of the Earth, Consideration of the movements, of continental plates, isolation, and evolution of temperate and tropical forests of Australia’s flora by Barlow and Hyland (1988) provide insights to the complex paleogeographic and paleoclimatic derivation of the remarkable diversity of this region, For nearly two centuries the causes for the striking differences in tree species diver- sity among regions of the Earth has interested biogeographers and ecologists. Wet tropical forests have 10 times as many trees species as moist forest of the northem and southern temperate zones at several spatial scales. Various explanations have been offered. Yet no ‘one mechanism has gained general acceptance, and several may contribute to the complex patterns. An outstanding review of regional tree diversity and comparison of temperate- zone diversity is presented by Latham and Ricklefs (1993b). They observe that ecologists ‘generally have invoked local-scale deterministic processes of competition and other inter- actions to explain global-scale diversity patterns. It is 2 large step from local-scale processes to such regional-scale predictions, Nevertheless, differences in diversity be- ‘tween regions with similar climate suggest that local and regional processes contribute separately to regional species richness. Their analyses indicate that contemporary patterns of tree species diversity owe much to historical and evolutionary events, as noted for the beeches. Latham and Ricklefs (1993b) provide a comprehensive analysis of present-day taxo- nomic diversity among major moist temperate forest regions of the Norther Hemisphere: (1) northern, central, and eastern Europe, (2) east-central Asia, (3) Pacific slope of North ‘America, and (4) eastern north America. A total of 1,166 species make up the characteris- tic tree flora of these regions, approximately in the ratio 2:12:1:4 (Table 20.5); eastern Asia has by far the most species and genera. ‘A comparison of genera that occur in either two or three of the regions (i.e. those that are neither endemic nor cosmopolitan ), is presented in Figure 20.11. Of these 63 genera, 59 (94 percent) occur in easter Asia. Regions sharing the most genera (20) are Table 20.5. Summary of Moist Temperate Forest Trees in the Northern Hemisphere by Taxonomic Level and Geographic Region. ‘Number of tre taxa characteristic of moist temperate forests in: Northern, Pacific central, & slopeof Eastern. Northern ‘Taxonomic Eastern == Eastcentral North North Hemisphere level Europe Asia America Ameriea (otal) gener Eee Pee Subclasses 5 6 9 10 Orders 16 14 26 39 Families 2 9 46 ” Genera 3 r 37 90 213 Species 124 ns 6 253 1,166 “Source: Aher Latham and Rickles, 19938, Reprinted with permission from Species Diversity in Ecological ‘Communities, Rickefs and Schluter, eds, © 1993 bythe University of Chicago. Reprinted with persion of the University of Chicago Press)Diversity at Continental and Subcontinemtal Scales 599 Paci easton \ North Noth) America ‘America Re Figure 20.11. Numbers of tree genera native to either two or three of the four north tem- Perate forest regions. Straight lines and the middle curved lines (bowing ut) indicate genera that occur in two regions. The inner curved lines (bowing in) indicate genera that occur in three regions including east central Asia. The outer ellipse indicates genera that occur in the three regions excluding east-central Asia. (After Latham and Ricklefs, 1993b. Reprinted with permission from Species Diversity in Ecological Commu- nities, Ricklefs and Schluter, eds., © 1993 by the University of Chicago, Reprinted with permission of the University of Chicago Press.) astern Asia and caster North America, reflecting the well-known range disjunction of ‘many plants inhabiting both regions (Boufford and Spongberg, 1983). Despite their prox- imity, the temperate forests of eastern and western North America share few genera and none uniquely. Eastem Asia emerges overwhelmingly as the core area of temperate tree genus distributions of the Northern Hemisphere. Why the high species richness in eastern Asia? The marked differences in diversity of temperate tree floras between east-central Asia and other regions of the Northern Hemi sphere is probably ancient (Latham and Ricklefs, 1993b). It appears to have arisen from differences between regions in colonization history and perhaps in subsequent rates of proliferation of endemic taxa. Colonization appears to have occurred more frequently in Asia, giving rise to new native taxa and their geographical spread among the temperate re- sions. The species proliferation in Asia is reflected in the occurrence of markedly more species at the local level than found in temperate forests elsewhere, Reasons for this in clude the enormous site differentiation provided by ubiquitous mountainous terrain and the lack of significant influence of continental glaciation, Current geographical distribu- tions and the fossil record suggest that most cosmopolitan taxa of temperate trees origi- nated in castem Asia and dispersed to Europe and North America. Furthermore, differ- ences between temperate and tropical flora reflect a physiological barrier to colonization of temperate zones that can be crossed only by the evolution of freezing tolerance mecha- nisms. Overall, explanations for latitudinal gradients in richness can be traced primarily to historical and evolutionary factors affecting land masses and floras rather than to present- day ecological interactions.600 chapter 20 / Diversity GLACIATION Continental glaciation has had marked effects on species diversity, and many of these ef- fects are described in Chapters 3 and 22. Extinctions have occurred in areas where species ‘were unable to migrate to their former locations. Elimination of species was severe in cen- tral Europe where the Alps acted as a barrier to southerly migration of species, The mas- sive continental ice sheets altered the climate of areas that were located at great distances from the advancing or retreating ice fronts. Such differences for eastern North America are discussed in Chapters 3 and 22 and seen in Figure 3.5. Glaciation has also affected sea levels around the world. Low sea levels during the Pleistocene Era caused formation of a land bridge between Asia and North America al- lowing immigration of humans into the Americas. The presence of humans has been asso- ciated with elimination of many of the large North American mammals such as the saber- tooth tiger and the American horse. As we have seen in Chapters 16 and 17 on disturbance and succession, human alteration of disturbance regimes of fire and flooding has been ‘widespread. The extent to which humans have influenced diversity in the Americas is un- known, but their introduction during the Pleistocene certainly had a remarkable effect on the structure and diversity of North American forests. LATITUDE AND ELEVATION Plant richness decreases with both increasing elevation and increasing latitude toward the Arctic Billings, 1973, 1995). Reasons for this decrease are very long-term evolutionary ‘and geographic processes and local ecological conditions such as decreases in mean an- nual and growing season temperatures, drought stress, and ultraviolet-B irradiation at high elevations (Billings, 1995). ‘The latitudinal gradient in diversity of many organisms from the tropics to the poles has generated great scientific interest. Wilson (1992) illustrates this gradient by numbers of breeding bird species by a slice through the northern hemisphere using land areas of roughly the same size: Greenland, 56; Labrador, 81; Newfoundland, 118, New York State, 195, Guatemala 469, and Columbia, 1,525. He notes that 30 percent of the Earth’s bird species occur in the Amazon Basin and another 16 percent in Indonesia. Tree species show an even more striking and ecologically-significant gradient of decreasing richness with increasing latitude, The remarkable tree and plant richness of the wet tropics is well known and is discussed in Chapter 22. However, the latitudinal relationship is complex due to marked differences in physiography and climate of regional ecosystems within any continent, For example, the geographical pattem of tree species richness for much of North America, illustrated in Figure 20.12, shows that factors other than latitude affect richness, ‘Tree richness is greatest in the southeastern United States and decreases steadily north- ward. It also decreases in the arid parts of the continental interior showing that latitude (ic., primarily temperature-related factors) is not the only determinant of richness. A strong relationship was found between tree richness and annual actual evapotranspiration and mean annual temperature when large geographic quadrats were used in computation (igure 20.12, Currie and Paquin, 1987). However, atthe fine scale, no significant relation- ship was found between actual evapotranspiration and local tree richness (0.5 to 10-ha plots) of broad-leaved deciduous trees (Latham and Ricklefs, 1993a,b). The tre richness pattern in North America reflects the evolutionary history of broad-leafed trees and the rel- ative newness of continental arctic climates compared to the unglaciated areas of southeast em United States (Latham and Ricklefs, 1993a,b) as well as local ecological interactions. lant diversity is strongly related to elevational gradients that are steeper up the sides of mountains than latitudinal gradients. Generally, plant diversity decreases as mountain,wh tude BA ation ition O-ha tain Divorsity at Continental and Subcontinental Seales GO Figure 20.12. Species richness of North American trees north of the Mexican border. Contours connect points with the same approximate number of species per quadrat. Data is based on 620 native tree species in quadrats each 2 1/2" x2 1/2" south of 50° N, and 2 1/2° x 5° (longitude) north of 50° N (Reproduced with permission from Nature (Currie, D. J., and V. Paquin, Large-scale biogeographical patterns of species richness), ©1987 ‘Macmillan Magazines Limited.) clevations increase (Billings, 1987). This relationship is especially true when elevation is ‘combined with increasing latitude as illustrated in Figure 20.13 for relatively high moun- tain ranges in Burope and Asia, ‘An excellent example contrasting patterns of plant species diversity in forests of two mountain ranges and two markedly different regional ecosystems of the western United States is presented in Figure 20.14, In the relatively humid, maritime climate of the ‘Siskiyou Mountains of southwestern Oregon, the slopes are forested at all elevations that were sampled. Richness of woody plant species decreases with increasing elevation; herb diversity is highest at middle elevations and lower in the dense stands with a sclerophyll602 Chapter 20 / Diversity oNzow vTaew i. SA IN 000” 5000" 600 Elevation n) Figure 20.13. Relationship of the number of plant species with increasing elevation in the European Alps (left) compared with high mountain ranges in central ‘Asia (tight) having many peaks above 6,000 meters. The letters show the increasing number of species with decreasing latitude: A = Alps, P = Pamirs, H = Hindu Kush, T = Tibet, N = Nepal Himalaya. (From Breckle, 1974, Reprinted with permission by the Council for Nordic Publications in Botany, Copenhagen, Denmark.) tree stratum at low elevations (Whittaker, 1960). The combination of these two trends pro~ duces the pattern of maximum richness in mesic forests at middle clevations (ca 1,200 0), with a secondary maximum at lowest elevations. Notice that at all elevations, richness in- treases from xeric (ridges and SSW aspects) to mesic and wet ravines. In the drier, conti- ‘hental climate of the Colorado Front Range of the Rocky Mountains, the forests are more ‘open and without sclerophyllous tees. Forests give way to shrubland and grassland at low Clevations and to alpine meadows at high elevations. Plant richness is low in forests of middle elevations and middle topographic positions of exposed, open slopes. Richness is higher in ravine forests the transition to alpine meadows, and the more open stands of dri- csi topographic situations and low elevations. Again, richness increases from xeric aspects to mesic and wet ravines. “These examples emphasize the importance of considering biodiversity through stu ies of regional ecosystems (southwestern Oregon mountains versus Colorado Rocky Mountains) and the local ecosystems (mesic ravines, open mid-elevation slopes, xeric ridges, subalpine woodland etc.) that occur at positions along gradients of temperature ‘and moisture in the respective mountain systems. Diversity at Local Scales “There are few sure or “global” generalities or principles that we can offer about biodiver- sity, For example, fertile soil may or may not promote diversity, and biomass productivity may be positively or negatively correlated with diversity. However, two concepts seem reasonably clear. Heterogeneity promotes diversity at all scales, and extreme conditions tend to constrain diversity. For example, a heterogeneous pattern of ecosystems or their components (climate, physiography, soil, hydrology) and heterogeneous disturbance regimes and stand structures tend to promote diversity In contrast, the most extreme phys-Diversity at Local Scales 603 q Woy powdey ‘L161 OAEMIUA AY) “wontsod onder Te1anos 20} paSezaae svespenb a pads simmjd @ ouex MS 3s N sss MN salou sade ued poroNOUS_seuney Brrr trea Oreo SIN ofSeNY 19M é 3 id 180104 jon, 8 “Happs § euch S05 a Ing wna} 2mp Jo UONSStuDd EA KBoporg poswow OSeA 404 SIP SIOQUINN “o Jo aduvy uous a yo adoys wiayseo omp pur UoRaIE warsaMNNOs Jo suIEUMOPY NOKTYS! “pro ams ex MS N _ NN 980m "0m, sss mo aN s00Pi sodoiS uedo _paleyous_souney 200" oe a \ e6insiopnes - mMydbisos coz |ren\ eve 1 000° fe-02 sey 8zp_---~ te euEUON, . sisev04 outdegns euesuavow 6s zer_cesz zie siz Aysiong wea pue weueg 80103 ‘Buty juar4 opeso}09 ‘aug wo Aysionig wep pue oH 010 SUBIUNOW RODS604 chapter 20 / Diversity ical conditions of temperature, water, soil reaction, and combinations thereof tend to limit biodiversity. Above all, however, biodiversity is most closely related to the spatial pattern, diversity, and key characteristics (physiography, climate, soil, and disturbance regimes) of regional and local ecosystems. It is beyond the scope of this chapter to examine the enor ‘mous array of factors and multiple-factor determinants of species diversity so that only se lected ecological relationships are discussed in the following sections. PHYSIOGRAPHY AND SOIL. ‘At meso- and microecosystem scales, physiography and soil, as described in Chapters 10 and 11, strongly influence the distribution and diversity of plants. Physiography, in partic- ular, because of its elements of form and parent material markedly affects plant diversity by its effects on microclimate and soil development. Diversity in mountainous regions in the temperate zone and in unglaciated areas is often very high because of the dissection Ge, heterogeneity) of terrain and the diverse sites and niches available for plant occu- pancy. Figures 2.12, 10,2, 10.4, 10.12, 11.8, 13.11, 15.1-3, and 6; and 20.14 all illustrate the striking effect of physiography as it mediates diversity in microclimate and soil water and nutrients Microtopography increases species diversity by providing highly localized environ- ‘mental heterogeneity (Chapter 10, Figure 10.15; Foster, 1988a). Regeneration of some un derstory tolerant species depends on microtopography, either of mounds of treefalls or from coarse woody debris. For example, yellow birch and eastern hemlock seedlings es- tablish preferentially on moss-covered logs and mounds in hemlock-northern hardwood forests (Chapter 5) as do seedlings of western hemlock and sitka spruce in old-growth forests of the Pacific Northwest (Harmon, 1987; Harmon and Franklin, 1989). Soil conditions of water, nutrient, and O, availability account for many observed dif- ferences in diversity. Soils that are extreme in some attribute—low water or nutrient avail- ability, high in specific mineral content (e.g., soil derived from serpentine parent mater- jal), or very shallow—often support forests low in tree richness (¢.g., pine barrens, black spruce swamps). However, such soils may support many herbs or rare and unusual herbs that have adaptations giving them a competitive advantage. Soils that are high in nutrient availability may support high tree species richness. For example, for a broad range of ecosystems in the Piedmont Province of North Carolina, ‘vascular plant richness was strongly related to soil nutrient availability (Figure 20.15). High richness was also associated with abundant moisture, and among sites with low soil fertility, the moister sites had more species. Soils that are deep and also high in water and nutrient availability may support either high- and low-diversity forests depending on the history and amount of disturbance and successional stage. Dominance by one or a few species often occurs more rapidly under favorable conditions, where all organisms can potentially grow rapidly (Huston (1993, 1994), Under such conditions, the best competitors are able to eliminate most other species by monopolizing an essential resource such as light. Under poorer soil water and nutrient conditions, the superior competitors are not able to dominate as rapidly, and many species are likely to coexist, leading to individuals that are smaller in size, fewer in num. ber, or both. In mesic beech-sugar maple and hemlock-northern hardwood forests of the western Great Lakes region, canopy coverage in late-successional stands may be very dense: Rel tively few species survive in the understory and ground-cover layers other than young indi viduals of the canopy species (Figure 15.16). This relationship for a mesic hemlock-north- cern hardwood forest is illustrated in Figure 20.8. Ecosystems in this fertile and moist Colonial Point moraine exhibit relatively low ground-cover diversity (Table 20.4). How.Diversity at Local Scales 605 Mosic Eutrophic © . tui ony Eutrophic 1s Mesie Mesotrophio Dry Mesotrophic 40}, Dy Olgotroptic ‘Goot Monadnock 1 2 3 4 6 Soll Cation Concentration (meq/100g) ‘Species Richness (Specis in 0.1 ha) Figure 20.15. Relationship between species richness (average number of plant species in a 0.1-hectare sample) and average exchangeable cation (Ca, Mg, K) for ‘of ecosystem types, representing wet, mesic, and dry sites, of the North Carolina Piedmont. (After Peet and Christensen, 1980b. Reprinted with permission of the ETH Zirich, Finanzdienste, (CH8092 Zurich, Switzerland.) ever, recall from the discussion above that a bigtooth aspen community, with relatively- open crowns, on a similarly fertile and mesic soil (Table 20.4, interlobate moraine) has approximately 40 percent greater ground-cover richness due primarily to favorable light conditions. In this case, catastrophic fire replaced the northern hardwood forest with as- pens. Ground-cover diversity observed in summer in ecosystems with a closed canopy may be somewhat misleading because of a relatively diverse flora of early spring cephemerals that complete much of their aboveground growth before the canopy trees leaf- out in the spring, COMMUNITY COMPOSITION AND STRUCTURE ‘The diversity of birds, mammals, and other forest animals may be affected by: (1) the species composition of forest stands, and (2) community heterogeneity or structurc/age differences within and among communities, vertical layering of vegetation, and horizon- tal arrangement of trees and other vegetation within and among stands (e.g., random ver- sus patchy stem distribution). A classic case is the relationship of bird species richness and structural complexity of vegetation or foliage height diversity. Bird species diversity thas been shown to be positively correlated with foliage height diversity in northwestern North America (MacArthur and MacArthur, 1961) but negatively correlated with it in northern Patagonia, Argentina (Ralph, 1985; Huston, 1994, p. 41). The northern spotted ‘owl (Strix occidentalis), a rare inhabitant of old-growth forests of the Pacific Northwest, requires large hollow trees for nesting and open subcanopy areas for foraging. The spot- ted owl controversy, a watershed event of American environmental policy (Yaifee, 1994), revolves around conserving structurally-heterogencous old-growth forests of the Pacific606 chapter 20 / Diversity Northwest to retain habitat of the owl and many other organisms of these unique forests (Franklin et al,, 1981; Franklin et al., 1997; see Chapter 21). In contrast, the red-cockaded woodpecker (Picoides borealis), a rare bitd of southeastern United States, is associated with old-growth pine communities with low structural diversity. This woodpecker nests in colonies in cavities that are carved over generations in large, living, fungally infected, resin-rich, longleaf, slash, or loblolly pines (Lennartz, 1988; McFarlane, 1992). The birds preferentially reside in ecosystems having large trees with little or no mid-story, few tall shrubs, and generally sparse ground-cover. Protection from predators, hot fires, ‘and availability of insect prey appear associated with the bird’s habitat—one with simpli- fied vertical structure. DISTURBANCE AND SUCCESSION Itis difficult 10 generalize about the effect of disturbance on species diversity. Ecosystems are $0 diverse in composition, structure, and function that they defy generalized textbook examples, Disturbance, being ecosystem specific, typically maintains a diversity level of trees, ground-cover, and animals consistent with the geographic setting of regional ‘ecosystems and the characteristic physiography, soil, and vegetation of local ecosystems (see Chapters 16 and 21). Diversity patterns of hurricane-prone ecosystems are different than fire- or flood-prone ecosystems. Overall, many ecologists have found that low species diversity occurs at both very high and very low frequencies and the highest dives sity at intermediate frequencies of disturbance (Huston, 1994). This concept applied to tree species of a tropical rain forest in Uganda is illustrated in Figure 20.16. Depending on ecosystem type, disturbance can be perceived to maintain either high or low species diversity. For example, frequent fires in presettlement time maintained some dry, outwash-plain ecosystems in the Lake States with jack pine as the only domi- nant tree species, whereas tree species richness increases markedly in the adjacent mesic moraines in which fires are virtually absent. Simultaneously, periodic fire in the outwash- plain ecosystems maintains high herbaceous richness because the herbs die out as the jack pine canopy closes as the trees develop in crown density with ag ‘A universal example of fire disturbance reducing temporal species richness is ob- served in ecosystems where mesophytic, fire-sensitive species colonize the site during the period of fire absence. In these situations, richness increases, but then decreases at the oc- currence of the next fire. For example, in fire-prone ecosystem types in northern Florida and southern Georgia, diversity of canopy trees is low in the sandhill, flatwoods, and cy- press ecosystems, each being nearly monotypic (Monk, 1967). Diversity of saplings and seedlings is higher than that of canopy trees and includes hardwood specie’ that colonize the understory—only to be eliminated or reduced in size by the next fire. In contrast, the southern mixed hardwoods, mixed hardwood swamps, and bayheads that seldom burn, the same richness of species is found in both the canopy and the understory. In some ecosystems, especially those characterized by periodic fires, frequent, less severe disturbances are necessary for the maintenance of diversity. For example, lack of periodic fires in prairie or savanna ecosystems typically leads to the encroachment of woody plants (Curtis, 1959) and a reduction in herbaceous diversity. In an attempt to re~ store resettlement oak savanna structure and composition to @ northem pin oak commu- nity in central Minnesota, annual prescribed burning was begun in 1964 (White, 1983; 1986). After 13 years of burning, the basal area of oaks was significantly reduced in burned versus unburned plots, Shrub cover was eliminated in bumed plots, and the rich- ress of herbs significantly increased in burned plots by about 60 percent. Overall, fie is an extremely important factor affecting biodiversity because pyrodi versity promotes biodiversity (Martin and Sapsis, 1991), Pyrodiversity, the variety in inDiversty at Local Scales 607 DIVERSITY Low — DISTURBANCES FREQUENT ———> INFREQUENT DISTURBANCES SMALL > Lange ‘SOON AFTER DISTURBANCE ——» LONG AFTER, CANOPY UNDER. STORY A COLONZING —& MIXED © cuMaAx Figure 20.16. General relationship between tree species diversity and disturbance re- lated to frequency of disturbance, size of disturbed area, and temporal scale corresponding to carly- (colonizing), mid- (mixed), and late-succes- sional stages of succession in the wet tropical Budongo rain forest of Uganda. The diagram represents the “intermediate disturbance hypothe- sis” where a mixture of early and late-successional tree species exhibits the greatest tree-species diversity at an intermediate position in distur- bance frequency and size and time after disturbance. (After Connell, 1978. Reprinted with permission from Science 199, p 1303, © 1989 by the American Association for the Advancement of Science.) tervals between fires, seasonality, size, and fire characteristics, has produced biodiversity atall ecosystem levels and worldwide from prehistoric times onward. Disturbance markedly affects species diversity in successional sequences because these are ecosystem specific. Overall, total plant species diversity is closely associated With the four general stages of secondary succession following disturbance (Chapter 17; Figure 17.4): (1) stand initiation or establishment, (2) stem exclusion or thinning, (3), un- erstory reinitiation ot transition, and (2) old-growth, We may characterize diversity of vascular plants in layers for the respective stages as follows: (1) high diversity in an open site soon after disturbance as species resprout or colonize by seedlings, (2) low diversity as dense overstory may “shade-out” all but the most understory tolerant species, (3) low but increasing diversity as gaps appear in the overstory (Figure 17. 14), and (4) low to high diversity depending on canopy density and number of gaps, as minor and major dis- turbances affect the declining old-growth forest more than younger stands of earlier stages. Examples of this general pattern of diversity with successional stage are given for Rocky Mountain forests (Peet, 1978) and for old-field mixed conifer-hardwood forests of the North Carolina piedmont (Christensen and Peet, 1981; Peet and Christensen, 1987). Obviously, disturbances occurring during these arbitrary stages would affect diversity in different ways depending on ecosystem type and disturbance characteristics —kind (wind, fire insect, ec.) origin (natural or human), and severity608 chapter 20 / Diversity HUMAN EFFECTS AND CONCERNS FOR DIVERSITY ‘Species extinction has escalated with increasing human populations and domestication of natural ecosystems by intensive agriculture, forest management, and urbanization. Main- taining the local, regional, and global diversity of species has become an increasing focus of ecologists, managers, and the public. An entire science of conservation biology has arisen in response to a desire to maintain species richness. Of special concer is the man- ‘agement of species whose ranges or habitats are highly limited in area, conservation of species whose presence or absence greatly influences the ecosystems in which they re side, and reduction of invasive exotic species that have the potential to outcompete native species or alter ecosystems in which native species occur (see Chapters 14 and 19). In the following sections, we examine several of these topics. Endemics and Rare and Endangered Species Much of the concern about diversity arises around species that are considered rare, in dan- ger of extinetion, or endemic, Endemic species are those whose ranges are extremely lim- ited in size, being restricted to an ecological region, a single mountain range, serpentine soils or other spatially-limited sites, or to politically-bounded units such as states or provinces. Endemics occur in areas that may or may not be considered diverse. However, tendemies often contribute substantially to regional diversity Endemism is not uniform across North America. In the United States, California, ‘Texas, and Florida have the highest numbers of endemic species (Gentry, 1986). Not all endemies, of course, occur in forest ecosystems, but many do. The majority of Florida’s ‘endemic plants are associated with swamps, flatwoods, and hardwood forests in northern ‘and central Florida, A large number of species is associated with a spatially-limited area known as “serub” (Myers, 1990). Many plants with very limited distributions apparently arose on isolated islands during periods when most of the current peninsula was below sea level. These species persist today in isolated scrub “islands.” An array of animals also evolved on these islands, and they include the Florida scrub jay (Aphelocoma coerulescens var. coerulescens), a showy and sociable bird, and the remarkable sand skink (Neoseps reynoldsi) which “swims” just below the sand surface. While these species are well adapted to survive on very specific sites and on an archipelago of serub islands, they are also regionally rare, and many are considered to be endangered, due to encroachmer of agriculture and urbanization into the ecosystems that support them. Keystone Species What is a keystone? A keystone is the central, topmost stone of an arch, popularly thought of as holding the others in place, or more generally: that one of a number of assorted parts that supports or holds together the others (Neufeldt and Guralnik, 1988). This concept can’t be taken too literally, however—as did the writer who observed that if all the pines of a forest were wiped out, “the ecosystem would totally disintegrate.” But ecosystems are ‘more than species; ecosystems change, slowly or drastically, but they don’t “fall apart” for Joss of a particular species, Instead, our perception is that the pine community, which we once knew intimately, upon catastrophic change is gone (i.e., had “disintegrated”), and the new community is a major force in building a new ecosystem. Dominant species are not necessarily considered “keystone” species. In ecology, the definition of keystone species has been expanded from the original animal ecology usage of Paine (1969) to identify a keystone species as one whose impactlates of Forest Management on Diversity G09 on its community or ecosystem is large, and disproportionately large relative to its abun- dance Power et al,, 1996). cologists recognize that the most abundant species play major roles in controlling the rates and directions of many ecosystem processes. These dominants are often crucial for maintaining their communities because they typically pro- vide the major energy flow and the physical structure that supports and shelters other or zganisms (Gentry and Dodson, 1987; Ashton, 1992). An excellent example is old-growth Douglas-fir trees that dominate their community. They are significant in ecosystem func- tion (Franklin et al, 1981) and support multitudes of organisms on their boles and branches (Chapter 21, Figure 21.1). However, many experiments have shown that some less abundant species have major community and ecosystem effects. Thus keystone species differ from dominant species in that their effects are much larger than would be predicted from their abundance (Power eta, 1996). Identifying keystone species is diffi- cult because we have learned to think of dominant species as playing key roles in forest ecosystems. Nevertheless, identifying keystone species will lead to a better understanding of how loss of species will affect ecosystem function “The Rosetta stone for identifying keystones is provided by Power etal, (1996). Key- stone species are often, but not always of high trophic status. They can exert effects, not only through consumption (e.g., pocket gophers, elephants, wolves, predatory ants and trout, kangaroo ras, ete.) but also by such interactions and processes as competition, mu- tualism, dispersal, pollination, disease, and by modifying abiotic factors (Bond, 1993; Mills et al., 1993). Possible examples of ecosystem modifiers or “ecosystem engineers” (Lawton and Jones, 1995) include beavers that swamp-out meadows and forests and bad gers whose mounds maintain diversity in prairie floras (Platt, 1975). Few woody plants have been identified as keystone species by Power etal. (1996); fig tees (Ficus spp.) that provide food for animals was the only woodland keystone identified “The importance of the keystone concept is in helping identify the most suitable areas for biodiversity preserves and in understanding the complex linkages among ecosystem biota and site-biota interactions. Power et al. (1996) gave three useful insights from the keystone concept: (1) land managers should carefully consider the consequences of the Joss of species for which no obvious role in ecosystems has been discovered, (2) intro- duced alien species may, like keystone species, have potential strong effects dispropor: tionate to their biomass, and (3) thore is a lack of well-developed protocol of identifying keystone species; the field is littered with far too many untested ancedotal “keystone species” Effects of Forest Management on Diversity Silviculture and forest management include many operations, such as regenerating, tend- ing, and protecting forests (Smith et al., 1997). Traditional management procedures, how- cever, are typically very different from the processes occurring in natural stands. The ere- ation of even-aged, even-sized stands using clearcutting and artificial reforestation (Oliver and Larson, 1990; Smith et al., 1997) has become the most common managerial system. Such stands are highly simplified and lack many components of community structure, such as snags and logs, as well as stand-level structural complexity, such as multiple canopy layers, irregular tree spacing, and gaps (Franklin, 1995). Forest ecosystems may be simplified at several levels: genetic, individual stand, landscape, and temporal (succes- sion). Managed forests have been simplified in response to economic criteria—efficient management (e.g., whole-tree harvesting) and high productivity of commercially-impor- tant tree species—in the belief that much of the structural complexity found in natural610. chapter 20 / Diversity stands is not essential to sustained tree productivity of the site. Research on the impor- tance of stand structure and biological legacies as they affect ecosystem function, biodi- versity, and long-term site productivity are discussed by Franklin (1995, 1997) and Franklin et al, (1997) for Pacific Northwest forests (see Chapter 21). Clearcutting, because of the enormous changes to ecosystems and their biota, has been the subject of many studies, and a good review is provided by Boyle (1992). Diver- sity of animals and plants may increase or decrease depending on the specific sites, scale and patter of clearcuts, and the methods employed, including post-cutting burning or site preparation. The effect of clearcutting is typically not a simple reduction in species diver- sity, but rather a differential effect. Species dependent on old-growth or near-natural forests are often lost from the system, presumably until suitable environmental conditions are restored, as illustrated by the following example, "A detailed study of understory species diversity in natural and managed Douglas-fir forest of the Pacific Northwest, revealed that changes in understory diversity are fairly short-lived following clear-cut logging and stash burning (Halpern and Spies, 1995). Pop- blations of most vascular plants recover to original levels prior to canopy closure. How- fever, diversity may remain depressed for decades on severely burned sites. and some species may expetience local extinction. Silvicultural prescriptions that maintain or foster spatial and temporal diversity of stand structure and site conditions will be most effective in maintaining diversity, Practices associated with intensive, short-rotation plantation for- est, that preclude or delay the development of old-growth attributes, may result in long- term loss of diversity Fragmentation of forests by roads, agricultural fields, and clearcut patches also markedly influences biodiversity. The consequences of ecosystem fragmentation are not only the creation of isolated “islands” of various sizes but also include large changes in the physical environment that markedly influence species and gene pool diversity. The significance of fragmentation and edge effects have been considered by many ecologists and geneticists (Harris, 1984; Saunders et al., 1991, Ledig, 1992), and we examine it in Chapter 21 at the landscape-level. ‘Genetic diversity of forests (ie. species diversity and gene diversity within species) has been greatly influenced by human activities that affect evolutionary processes of ex- tinction, selection, drift, gene flow and mutation (Ledig, 1992), In some cases, diversity may be increased by exploitation and harvest practices (c.g. exposing recessive genes and increasing the occurrence of novelties and mutants), but these practices can also reduce diversity. Most forest harvest practices and associated road building cause fragmentation, result in drastic swings in population sizes, and simplify age structure—all of which affect the breeding systems of forest trees. Ledig (1992) describes how habitat alteration, envi ronmental deterioration, and domestication of forest species may all lead to reduced bio- tdiversity and, therefore, the urgent need to inventory genetic diversity of forest species ‘and monitor their changes. CONSERVING ECOSYSTEM AND BIOLOGICAL DIVERSITY 'As landscape ecosystems have become increasingly dominated by humans, the list of species considered to be in danger of extinction has grown (Vitousek et al, 1997). Cur- rently, more than 950 species or subspecies are considered to be in peril of extinction in the United States by the year 2,000 (Wilson, 1992). However, the concem is deeper than rare and endangered species; itis for the immense number of unknown species and for ecosystem processes that are basic to long-term sustainability of ecosystems. Endangered