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    Biodiversity Chapter 20 Part 1

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    CHAPTER 20 DIVERSITY Tu: Barues BV etal (eds) Forest Belegy » 4 Ect. “Vatu Weg RGous. New Pevk., ‘The diversity of organisms, measurement of diversity, and hypotheses about the causes of diversity have long interested ecologists. Exploring natural forests, making lists of species, hunting rare species, and posing questions such as why a species is present, rare, or abundant at some sites but not at others, is fundamental to the forest ecologist. Diver- is simple concept; in ecology it means variety. However, diversity grows rapidly in implexity with attempts to describe and measure it and compare it from one area to an- other. At its simplest level, the study of diversity is a descriptive pursuit involving count ing of listing species, ‘The study of diversity may also involve a range of foci from genes to ecosystems. It becomes closely intertwined with theoretical and practical questions related to ecosystem productivity and conservation. Numerous questions arise: Is species diversity a product of ‘site and its physical attributes? Is it a product of biotic interactions of competition and ‘mutualism within the community residing on that site? Are ecosystems with many species ‘more or less stable than those with only a few? How do we best identify and preserve the vatiety of organisms and ecosystems that exist locally, regionally, and globally? How do we manage forests in a manner that maintains their diversity? We have many good in- sights to these questions, but clearly we know too little about the genetic, species, and ecosystem aspects of diversity and their causes. ‘This chapter is organized into several sections. First, we consider concepts and val- tues of species and ecosystem diversity. Second, we examine the measurement of diversi and provide examples at species and ecosystem levels, Then we discuss controls and pat- tems of forest diversity, beginning first at a continental scale and progressively focusing ‘on more local landscapes. Finally, we discuss important issues related to management and conservation. While our treatment is primarily at organism and ecosystem levels, itis im- portant to recognize throughout thet, ultimately, diversity is closely intertwined with many 377 B78. Chapter 20 / Diversity interacting factors, including paleogeology and ecology, ecosystem geography, plant physiology, and human activities, Each chapter of this book can be thought of as present- ing one or more aspects of the physical and ecological processes whose ultimate result in cludes the diversity of ecosystems and organisms that inhabit them. The scope of our overview of diversity is necessarily limited, and readers are encouraged to delve deeply {nto books that treat organismal diversity (Wilson, 1992; Ricklefs and Schluter, 1993; Huston, 1994), measurement of species diversity (Peet, 1974; Pielou, 1975; Magurran, 1988: Krebs, 1989), and conservation biology (Soulé, 1986; Primack, 1993; Meffe and Carroll, 1997). CONCEPTS OF BIOLOGICAL AND ECOSYSTEM DIVERSITY Diversity can be examined from many different viewpoints. For example, the diversity of physical features of bedrock, landform, geomorphic processes, parent material, and cl imate is of ultimate importance to the distribution and growth of plants. The diversity of the biota—plants and animals, their gene pools and their arrangement in aggregates of populations and communities is of great interest and concern. Because biota are funda: ‘mentally different from the physical environmental factors and the ecosystems that sup: port them, we define biological diversity (biodiversity) as the kinds and numbers of or- ‘anisms and their patterns of distribution. This definition also includes properties of biota, including the genetic diversity of their gene pools and other types of diversity related to groupings of biota, such as the diversity of families and genera, populations and commu nities, and “structural” properties of communities (vertical layering, plant density, or patchiness, etc). ‘Although it is popular today to include landscapes and ecosystems as part of “biodi versity,” biota are not ecosystems and biodiversity should be distinguished from evosys tem diversity. Because biodiversity depends on ecodiversity (Rowe, 1992b, 1997), the di versity of ecosystems deserves separate and special consideration. Ecosystem diversity is defined as the kind and number of ecosystems in an area and includes the patterns of ass0- ciation of ecosystems with one another and the recurrence of these patterns in a given landscape. Ecosystem diversity can be estimated at multiple spatial scales by using the same diversity indices that are used to measure species diversity. Because species are inte {gral parts of ecosystems, understanding ecosystem diversity is fundamental (0 maintain. ing and conserving biodiversity ND ITS VALUE ;PECIES DIVERST The statement by Solbrig (1991) that scientists have a very rudimentary knowledge of biodiversity accurately characterizes our understanding of the Earth's biota. OF the 10 to 100 million species that are estimated to exist, only about 1.4 million species of plants ‘animals, and microorganisms have been described (Wilson, 1992). The breakdown of the organisms known to exist (Figure 20.1) shows that insects alone comprise 53 percent of the total, In the plant kingdom, angiosperms have been the dominant group for the past 150 million years and constitute about 18 percent of the total with dicots the dominant group by far (Figure 20.2). The immense combined diversity of insects and flowering plants is no accident because these groups are united by intricate symbioses (Wilson, 1992). The insects live on and in plants in their every nook and cranny, consume every anatomical part, and, as we recall from Chapter 14, a large fraction of plants depend on insects for pollination and dispersal. Despite this seemingly large number of organisms, virtually all plant and animal ‘Species Diversity and its Vaue 579 other animals _/ 281,000 Higher / Pants Viruses 248,400 Ae ns \ ior 30,800 \ (bacteria and similar forms) 4,800, OD PP Figure 20.1, Number of living species of all kinds of organisms currently known. The total number of all species is 1,413,000. Organisms are shown by major group. Insects and higher plants dominate the diversity of living organisms known to date, but vast arrays of species remain to be discovered in bacte- ria, fungi, and other poorly studied groups. Most flowering plants live on the land; algae prevail in the sea. (After Wilson, 1992. Reprinted by per- mission of the publisher from THE DIVERSITY OF LIFE by E. O. Wil- son, Cambridge, Mass.: Harvard University Press, Copyright © 1992 by the President and Fellows of Harvard College.) species that have ever lived on the Earth are extinct (Raup, 1986). Up to four billion species of plants and animals are estimated to have lived at some time in the geologic past (Simpson, 1952), yet there are far fewer living today. Thus extinction of species has been almost as common as origination. Major extinction events are regularly spaced in geo- logic time (Raup, 1986). There are several major mass extinctions, like that at the end of the Cretaceous, but relatively sudden and rapid turnovers occur at lesser scales as well For the past 600 million years, the course of biodiversity, despite extinctions, has been up- ward (Figure 20.3). Global biodiversity of plants Figure 20.4) reached its peak in the Cenozoic by: (1) creation of the aerobic environment, (2) fragmentation of land masses, and (3) increased packing of species into regional and local ecosystems of the developing landscape (Wilson, 1992). Note in Figure 20.4 that the number of plant species in local floras has more then tripled in the past 100 million years. Although this brief overview il- lustrates the immense increase in biodiversity, Wilson (1992, p. 343) observes that: “The 580 chopter 20 / Diversity Monocots 50,000 Minor Groups 1900) SB Gymnespoeme \ Bryophytes 529 16/600 130 tr = ( \ Figure 20.2. Number of living species of higher plants currently known. Total number of higher plant species is 248,000. Species are shown by major group, Plant diversity of the world consists primarily of angiosperms (flowering plants), which in turn make up grasses and other monocots and also a huge variety of dicots, from magnolias to asters and roses. (After Wilson, 1992. Reprinted by permission of the publisher from THE DIVERSITY OF LIFE by E. 0. Wilson, Cambridge, Mass.: Harvard University Press, Copyright © 1992 by the President and Fellows of Harvard College.) sixth great extinction spasm of geological time is upon us, grace of mankind, Earth has at last acquired a force that can break the crucible of biodiversity.” Therefore, itis useful next to consider the value of biodiversity; later in this chapter we consider the increasin role of biodiversity in shaping the science of ecosystem conservation, management, and restoration Value of Biodiversity The reasons for conserving, promoting, and managing a rich variety of organisms are ‘many (Burton et al. 1992). These include: The intrinsic value of the existence of life forms and ecosystems (Leopold, 1949; Regan, 1981; Norton, 1982; Naess, 1986; Rowe, 1990; Mosquin etal, 1994) Aesthetic or anthropocentric values; nature preserves, wilderness, and biological di- versity can all play a role in promoting human well-being (Kaplan and Kaplan, 1989; Easley et al., 1990; Kaplan, 1992; Thompson and Barton, 1994) Number of Famiies Figure 20.3. 3 5 z < SILURIAN NAN PERMIAN TRIASSIC “JURASSIC ‘ORDOVICIAN DEVO! (CRETACEOUS “CARBONIFEROUS — Nllions of Years Ago Example of the increase of biological diversity over geological time using data from families of marine organisms. A slow increase is seen, with oc~ ‘asional setback through mass global extinctions. There have been five such extinctions so far, indicated here by lightning flashes. A sixth major Gecline is now underway as the result of human activity. (After Wilson, 1992. Reprinted by permission of the publisher from THE DIVERSITY OF LIFE by E. . Wilson, Cambridge, Mass.: Harvard University Press, Copyright © 1992 by the President and Fellows of Harvard College.) Local Diversity of Plants /ON.| CARBON. | PERM. [TRAS| JURASSIC | CRETACEOUS TERTIARY ce PALEOZOIC (in part) MESOZOIC ‘ENOZOIC | 408 Figure 20.4. 360 320 288 240 213 148 6 76 Milfons of Years Ago ‘The average number of plant species found in local floras over geologic time. The number of plants found in local floras has risen steadily since the invasion of the land by plants 400 million years ago. The increase reflects @ growing complexity in terrestrial ecosystems around the world. (After Wil- fon, 1992. Reprinted by permission of the publisher from THE DIVER- SITY OF LIFE by E. O. Wilson, Cambridge, Mass.: Harvard University Press, Copyright © 1992 by the President and Fellows of Harvard Col: lege.) 581 582 Chapter 20 / Diversity Important products other than fiber may be derived from forest organisms, e.g., wild species for food and medicine (McNeely, 1989). Certain species indicate the integrity of ecosystem processes, whereas others act as ‘warnings of critical stress thresholds of pesticides or atmospheric pollution that may endanger ecosystem function (Burton et al, 1992) Many species are important ecological indicators of site productivity (Chapter 13). Diverse organisms, and especially their gene pools, provide a sound resource base a5 the best insurance for coping with a rapidly changing and uncertain future environ- ment (Burton etal, 1992). Many species play mutualistic roles in the re ecosystems. Many species play key rol bance events. sneration and function of forest in the resiliency of ecosystems following normal distur ‘The importance of plant species for a variety of uses besides timber or fiber has been widely emphasized. Opportunities for identifying natural products for future pharmaceuti- cal, agricultural, or industrial value increase with species diversity. Pharmaceutical com- pounds derived from wild plants ate often of global importance. The Madagascar periwin. kle (Catharanthus roseus), for example, is the source of at least 60 alkaloids that are used in the treatment of childhood leukemia and Hodgkin's disease, and were reported worth ‘about USS160 million in sales each year (Shiva, 1990). Another medicinal example is the Wester yew tree, the source of taxol, a compound found to have strong activity against a umber of cancers (Wani et al., 1971), Wild species are not only used for food, medicine, and crafts in subsistence societies, but some are brought into cultivation such as was done ‘with kiwi fruit (Actinidia deliciosa). As Burton et al. (1992) point out, the valuable use of this and other species could not have been anticipated decades ago. A number of equally useful species are doubtless being destroyed today by short-sighted land management. ‘The evolution of these species and their useful attributes is, in part, a function of their membership in diverse and complex ecosystems. In addition to many specific values for biota already discussed, Costanza et al (1997) have estimated the market value of the world’s ecosystem services (ecosystem processes and functions such as food production, waste treatment, pollination, and re fation) to human welfare to average about US$33 willion (10!2). Global gross national product total is around USS18 trillion, Forests contribute 14 percent of the $33 tillior and wetlands (in part forested swamps and floodplains) contribute another 15 percent This bottom line approach is striking and provides a powerful incentive for conservation ff the natural capital stock that produces these services, Many ecologists and others would additionally, or alternatively, cite very real and “priceless” ethical and cultural rea sons for sustaining the Earth’s ecosystems. MEASURING DIVERSITY Levels of Diversity Diversity encompasses the two different concepts of variety: richness and evenness. Rich ness refers to the number of units (alleles, species, families, communities, ecosystems) per tunit area, and evenness refers to their abundance, dominance, or spatial distribution, These two concepts can be applied and measurements made at many levels of organization: Ecos- phere and continental-level ecosystems (macroecosystems, see Chapter 2), regional or Levels of Dwersity 583 mesoecosystems, local oF microecosystems, species, and the genetic diversity of allele fre- {quencies within populations of a species. The focus of biodiversity measurement is typi- cally the species, because species are easily observed and the most common taxonomic unit used in ecological studies of forest ecosystems. Species diversity can refer to all organisms in a community, but usually discussions are restricted on practical grounds to certain groups of species related to the interest of the ecologist. A single forest ecosystem type may contain many trees and shrubs and a great amber of herbs, bryophytes, birds, mammals, insects, and microorganisms. However, most studies ignore whole groups of organisms, especially those that are difficult to lo- cate, let alone to count or measure. Thus when studying diversity or comparing it between, ecosystems or regions, itis important to identify explicitly the set of organisms being com: sidered and the site-specific area of their occurrence. This point is especially important because forest ecosystems are remarkably different and complex; the diversity pattern of ttees may not reflect that of ground-cover vegetation, birds, or microorganisms. ‘Most studies of biodiversity occur at the local level in areas of <1 to ca 25 ha (micro- ecosystem scale, Chapter 2). Local site factors of microclimate, landform, and soil, as well as interspecific interactions determine the observed patterns of richness and even- ness. Diversity at the broad meso- and macroecosystem scales is sometimes referred to as landscape, or regional diversity in the literature. Here the focus may be on relatively broad vegetative cover types such as those in Yellowstone National Park (Romme, 1982; see Chapter 21) or the White Mountains of New Hampshire (Reiners and Lang, 1979). The pattern of diversity observed in such cases results from the superimposition of multiple patterns of: (1) species distribution along gradients of limiting factors, (2) recurrent spatial patterns of local ecosystems, and (3) communities in different stages of succession fol owing disturbance. In continental-level studies (Currie and Paquin, 1987; Latham and Ricklefs, 1993a,b), diversity of taxonomic units above the species level (.c., genera, fam ily, order) are often examined. At this scale, diversity is related to macroclimate, physiog- raphy, continental drift, and the historical development of plant floras. ‘Whittaker (1960, 1972, 1975, 1977) was one of the first to recognize that species di versity needs to be measured in several ways and at several scales. First, he recognized in- ventory diversity that could be estimated at four increasingly larger spatial scales: Point diversity: at a microsite scale with samples taken from within a homogeneous site, ie., microsite within a local landscape ecosystem type (¢.g., 100 t0 500 m?) Alpha diversity; at a homogeneous site, ie., for a given landscape ecosystem type or community in a homogeneous site (microecosystem level, e.g., < 1 to several hundred hectares; Table 2.4). Gamma diversity: at ¢ larger landscape unit, ie., for a regional ecosystem at the mesoecosystem level, ic.,a group of alpha-level ecosystems within a region (e.g, 62: 0 2,500 ha; Table 2.4), Epsilon diversity: total diversity for an area encompassing a group of areas of gamma diversity, ie., macroecosystem level (e. g., > 2,500 ha). Whittaker also recognized that species composition changes across the landscape, both along environmental gradients, and from one generally homogeneous site to the next. This change from one area to the next, regardless of scale, he referred to as differentiation di- versity or beta diversity, Besides community composition, diversity can also be used to describe the “struc turat” diversity of communities. In forested systems, this analysis focuses on the spatial pattem of species occurrence (random, systematic, contagious or patchy), stand density 384 chapter 20 / Diversity and diameter-class distribution, and the vertical layering of a forest from the tallest tree to the shortest plant on the forest floor, For example, Franklin et al. (1981) used a diversity index to confirm differences in spatial heterogencity (between-tree spacing intervals) that hhad been observed between old-growth Douglas-fir forests (450-year-old trees) and young-growth stands (25-year-old trees). ‘Structural diversity of communities is of considerable importance in wildlife ecology where the number of animals capable of making use of a forested system may depend heavily on the presence’ of appropriate areas for nesting, feeding, resting, and hiding ‘Structural diversity for old-growth, mixed-age or size, and mixed-species stands is typi cally high, whereas that for plantations or single-age, single-species stands is low. For ex. ample, the Kirtland’s warbler, a rare and endangered neotropical migrant nesting in ecosystems dominated by jack pine in northem Lower Michigan, prefers the patchy distri- bation of jack pine individuals following fire to the systematic arrangement of jack pines in plantations (see Chapter 21), ‘Measurement INVENTORY DIVERSITY: ALPHA DIVERSITY ‘There are various ways to measure the “diversity” of species. Whittaker (1977) described inventory diversity as having two components, richness and equability (evenness), and, as sve have described, cited four levels of scale, each reflecting successively larger land areas varying from a microsite to the macroecosystem level. “The simplest way to measure species diversity is to count the number of species pre- sent in a designated area. This number, or species richness, is the oldest, most fundamer tal, und perhaps the least ambiguous of the diversity measurements (Peet, 1974). It d pends on the area sampled, s0 richness is often expressed as number of species per unit fea, Richness varies with the size of the area sampled. In an area of homogeneous com- position, such as a single ecosystem type with a relatively little microtopographic varia tion, relatively small gaps or disturbance features, and uniform climate, richness will in- crease with increasing size of sampled area up to a point where all, or almost all, species have been captured in the sample, At this point, increasing the sample size results in no in ‘crease in species number. In determining the richness for a given ecosystem by measuring sample plots within it, it is important to use a plot size large enough, or enough plots, to encompass most ofthe species. In any sample of a large forested area, this total number is very dificult to determine for most major groups of organisms. Further, as the sampled area expands beyond a single local ecosystem, environmental heterogeneity increases, and different ecosystems with different species arc included in the sample. Therefore, to ob- tain a reliable estimate of alpha diversity of species for a given ecosystem, it is important to recognize that ecosystem's boundaries. Diversity also depends on the distribution of species within an area, termed evenmess cor equability, in other words, the degree to which all species share dominance in an area, For example, forest stands A and B in Figure 20.5 both have eight species; therefore, they have the same richness. However, stand A is clearly dominated by one or two species, with the other species being relatively uncommon. Stand B exhibits a more equal abun- ddance of all species and thus exhibits greater evenness. Which stand is more “diverse?” Stand B—because with the same number of species, it has the more equal abundance of the species. Note that each of the two diversity indices shown in Figure 20.5 has a higher value for stand B than for stand A. "The two diversity indices illustrated in Figure 20.5 combine richness and evenness in ‘one index value. This dual-concept diversity was introduced into the ecological liverature by Simpson (1949), and although generally accepted, it can be @ source of confusion (Peet, Measurement 585 1974), Most workers have considered that diversity should include both an evenness and a richness component. Nevertheless, as Peet (1974) observed, retention of diversity as a broad term encompassing all of the subordinate concepts, with a different terminology for cob o 49 17 2 28 20 G8 a7 a a5 49 «G87 GF 65 8D Dominance rank Figure 20.6. Comparison of ground-cover species diversity of two ecosystems using plots of species-abundance. a, ecosystem type 1 occurs on an outwash plain landform with excessively drained, infertile, sandy soil; b, type 116 ‘occurs on a moraine with moderately well fertile, loamy soil. Abundance is ‘measured by percentage cover. Ground-cover species for each ecosystem are ranked in order by their percent coverage along the x-axis (I=species with greatest abundance), Richness is indicated by the number of species, shown at the far right on the x-axis. Evenness is indicated by the shape of the curve, with ecosystem 116 being more “even” in distribution than ecosystem type 1, (After Pearsall, 1995.) short overstory of small bigtooth and trembling aspens (ecosystem type 1), and (2) a mod rately well drained, loamy, fertile moraine supporting a moderately dense canopy of tall bigtooth and trembling aspens, white ash, paper birch, and basswood (ecosystem type 116). In this example, percent coverage is used as the measure of species abundance. The species-abundance plot for type I shows lower ground-cover species richness (37 versus 69 species, Figure 20.6) and a sharp drop in dominance from the most dominant species (bracken fern, mean percentage cover: 44) to the second most dominant species (mean percentage cover of about 2). Such a plot is often observed in ecosystems that are extreme fn some way, such as in microclimate, fire frequency and severity, light level, or fertility, ‘and in an early successional stage. In contrast, the plot for type 116 illustrates nearly twice the species richness as type 1. It exhibits a much more gradual drop in dominance, from about 3.3 percent to <1 percent, from the most 0 least dominant species. This plot is more 588 chapter 20 / Diversity typical of ecosystems that are not at the extremes of microclimate, fertility, light, or dis- turbance, and in mid-successional stage, DIFFERENTIATION OR BETA DIVERSITY Besides species diversity within a given area, one may measure how different or similar a range of samples is in terms of the variety (and sometimes the abundance) of species found in them (Magurran, 1988). For example, communities of a given area that have very few species in common, collectively have a high beta diversity, and vice versa, This change or difference in species variety may be determined for ecosystems or communities ‘that occur along an environmental gradient or within an area supporting diverse sites. The often-used term beta diversity gives litte clue to the spatial or ecological context of the di versity that is measured. Thus beta diversity may be usefully understood as differentiation diversity (Whittaker, 1977; Magurran, 1988)—a measure of species diversity along a gra

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