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Does biodiversity play a significant role in ecosystem function? ASB Sustainable Land Use Mosaic (SLUM) Working Group Symposium 12-15 November 2001 Chiang Mai, Thailand CBM Definitions Biodiversity Conceptual: The variety of life on earth expressed in terms of gene, species and ecosystem. (cf. Heywood & Baste, 1996) Operational: The quantity and composition of species and functional types recordable in any area. (Gillison, 2001). Functional types: (FTs) are sets of organisms showing similar responses to environmental conditions and having similar effects on the dominant ecosystem processes. (Diaz, 1998). ASB SLUM Mtg Chiang Mai 11 Nov 01 Diversity …The number of different items and their relative frequency. For biological diversity these items are organized at many levels …. Thus the term biodiversity encompasses different ecosystems, species, genes and their relative abundance. (US Congress, Office of Technology Assessment, 1987). ASB SLUM Mtg Chiang Mai 11 Nov 01 Indicators and surrogates The vast number of biota that influence ecosystem function are mainly small organisms that are very difficult to measure. For most practical purposes their diversity and that of other larger, difficult-to-measure organisms, is assumed to be indicated by more readily observable units such as plant species and functional types. (This tends to be an act of faith commonly applied in private and denied in public). ASB SLUM Mtg Chiang Mai 11 Nov 01 Ecosystem 1. A community of interdependent organisms together with the environment which they inhabit and with which they interact. (Allaby, 1977). 2. A functional system which includes the organisms of a natural community together with their environment. (Lapedes, 1976) 3. … the sum total of vegetation, animal, and physical environment in whatever size segment of the world is chosen for study. (Fosberg, 1967) ASB SLUM Mtg Chiang Mai 11 Nov 01 The ecosystem concept is used widely but ambiguously. It can: a) contain only a functional meaning, b) have a spatial connotation which includes any level of scale or c) the spatial aspect can be included, but additionally, relative homogeneity must characterize the system (Johnson and French, 1981, quoted by Godron & Forman, 1983) ASB SLUM Mtg Chiang Mai 11 Nov 01 Differences between ecosystem and landscape: Ecosystems: are relatively homogeneous Landscapes: are relatively heterogeneous. A landscape is “…a kilometers wide area where a cluster of interacting stands or ecosystems is repeated in similar form” (Godron & Forman, 1983) ASB SLUM Mtg Chiang Mai 11 Nov 01 ‘Ecosystem’ conversion to ‘landscape’ ASB SLUM Mtg Chiang Mai 11 Nov 01 Landscape assemblages In NW Mato Grosso, Western Amazon basin Brazil ASB SLUM Mtg Chiang Mai 11 Nov 01 Species Elevation (m) 500 700 900 1100 1300 1500 1700 1900 2100 2300 2500 Plants Dipterocarpus tuberculatus Shorea obtusa Castanopsis sp. Chromolaena odorata Imperata cylindrica Smilax sp. Melastoma malabathrica Arisaema sp. Birds Collared Falconet Sooty-headed Bulbul Red Jungle Fowl Scarlet Minivet Striped Tit-babbler Grey-throated Babbler Arctic Warbler Range (elevational) distributions of some key taxa across different landscapes – Mae Chaem, Northern Thailand ASB SLUM Mtg Chiang Mai 11 Nov 01 Differing views of species role in ecosystems 1. Systematists and population biologists: – No two species can exist in exactly the same habitat – Every species has a unique resource requirement and hence use pattern ASB SLUM Mtg Chiang Mai 11 Nov 01 2. Ecologists: – More than one species can occupy a functional type or resource niche (e.g. mesophyte, xerophyte, producer, consumer, life form, C3, C4 path, nitrogen fixer, PFT…) – More than one functional type can occur within a species. Approaches to species, biodiversity and ecosystem function: Traditional: Community ecology: Species diversity is a dependent variable controlled by abiotic conditions and ecosystem constraints. Ecosystem ecology: Dominant species control ecosystem properties. Recent: Now consider role of biodiversity as a potential modulator of ecosystem processes. ASB SLUM Mtg Chiang Mai 11 Nov 01 Some general assertions • Ecosystem stability and primary productivity vary directly with diversity in species and functional type • Species properties expressed as functional types exert a greater degree of control on ecosystem function than species diversity (richness) ASB SLUM Mtg Chiang Mai 11 Nov 01 (After Springett, 1976) See also Swift & Anderson (1993) ASB SLUM Mtg Chiang Mai 11 Nov 01 Biodiversity and ecosystem function: Recent consensus* • Some minimum number of species is essential for maintaining ecosystem function under constant conditions • A larger number of species is probably required for maintaining ecosystems in changing environments • Determining which species have significant impact on which processes in which ecosystem remains an open emiprical question *Loreau et al. (2001) ASB SLUM Mtg Chiang Mai 11 Nov 01 Cont.. • Increased primary production via higher plant diversity can be expected to stimulate secondary productivity. • Changes in one trophic level may lead to a variety of potential responses for processes at higher trophic levels. • Mechanisms for generating primary productivity may range from systems with a few dominant species or functional types (low diversity) to systems with high diversity, low level dominance and high complementarity (synergy). ASB SLUM Mtg Chiang Mai 11 Nov 01 Hypothetical mechanisms involved in biodiversity experiments using synthetic communities Loreau et al. (26 October, 2001) Science: 294:804-808 Dual hypothetical mechanisms for species, FT diversity and ecosystem productivity Diversity of Spp and FTs Dominance Complementarity Low High (few groups) low High Low (many ‘ rare’ ) High Primary productivity Local, regional, random processes. Within and between landscape heterogeneity high ASB SLUM Mtg Chiang Mai 11 Nov 01 ? Local, deterministic processes. Niche diff’n high . Within landscape heterogeneity low Ecosystem functions and processes related to the transfer of energy, nutrients, water and genetic information Function/ Process Nutrient capture Decomposition and soil formation Mycorrhizal activity Photosynthesis Herbivory Pollination Species interactions (mutualisms, symbiosis, predation, parasitism, competition) Water uptake and loss Source: Hobbs (1992) See also Giller et al. ( 1997 ) for below-ground ASB SLUM Mtg Chiang Mai 11 Nov 01 Action Nutrient cycling Nutrient cycling Enhanced nutrient uptake Energy capture and productivity Energy and nutrient capture Genetic information transfer Energy, water and nutrient transfer Water transfer Plants as basal ecosystem units • Most terrestrial (mobile, heterotrophic) biota ultimately depend on plants for survival • Plants as mostly sessile, autotrophic units are readily observable in nature • Variation in plant form and function is measureable along key environmental gradients ASB SLUM Mtg Chiang Mai 11 Nov 01 Water Light (energy) High ASB SLUM Mtg Chiang Mai 11 Nov 01 Medium Low ASB SLUM Mtg Chiang Mai 11 Nov 01 Plant Functional Types* • Photosynthetic envelope (leaf size, inclination, chlorotype, morphotype) • Physical support system or life form based on position of perennating buds • Above-ground rooting system • Minimum set of attributes • ‘Coherent’ functional model * after Gillison (1981) ASB SLUM Mtg Chiang Mai 11 Nov 01 VegClass, Windows-based, user-friendly software for data entry and meta-analysis; integrated with field proforma to support rapid vegetation survey Plant Functional Types: leaf + stem photosynthesis, incl. parasites, carnivores ASB SLUM Mtg Chiang Mai 11 Nov 01 Ecosystem functions and processes related to the transfer of energy, nutrients, water and genetic information* Function/ Process Nutrient capture Decomposition and soil formation Mycorrhizal activity Photosynthesis Herbivory Pollination Species interactions (mutualisms, symbiosis, predation, parasitism, competition) Water uptake and loss * PFT relevance highlighted in red ASB SLUM Mtg Chiang Mai 11 Nov 01 Action Nutrient cycling Nutrient cycling Enhanced nutrient uptake Energy capture and productivity Energy and nutrient capture Genetic information transfer Energy, water and nutrient transfer Water transfer Patterns of richness in plant species and functional types under different land use types Indonesia : Jambi - Lampung Legend Cassava Imperata Mono. Plantation Natural Forest Logged Forest Agroforestry 160 140 120 Species 100 80 60 40 20 0 0 10 20 30 40 50 Modi Plant FunctionalTypes (modi) ASB SLUM Mtg Chiang Mai 11 Nov 01 60 Regression plot of termite abundance & basal area of woody plants [r2=0.985] over 7 land use systems: Jambi BS Nov. 97 100 Primary Forest Termite Abundance 90 Jungle Rubber 80 70 Secondary Forest 60 Rubber Plantation 50 40 30 Paraserianthes Plantation 20 10 Imperata 0 Cassava 0 2 4 6 8 10 12 14 16 18 20 22 24 Basal Area m2 ha-1 ASB SLUM Mtg Chiang Mai 11 Nov 01 Ratio of plant species richness to plant functional types as an indicator of Termites species richness [R-Sq = 0.97] over 7 land use systems : Jambi BS Nov. 97 4 0 Termite species richness Primary Forest 3 0 Secondary Forest Jungle Rubber 2 0 Paraserianthes 1 0 Plantation Rubber Plantation Imperata 0 Cassava 1 . 0 1 . 2 1 . 4 1 . 6 1 . 8 2 . 0 2 . 2 2 . 4 2 . 6 2 . 8 3 . 0 Ratio of plant species to plant functional types (modi) ASB SLUM Mtg Chiang Mai 11 Nov 01 Correlations between key plant variables, fauna and above-ground Carbon Jambi Baseline survey Attribute Species Modi Spp/Modi Ground-dwelling Termite abundance 0.872 0.766 0.946 Termite species 0.849 0.698 0.976 Lep/ground 0.834 0.790 0.920 Canopy: Unident. insects 0.771 0.418 0.839 Collembola 0.643 0.089 0.882 Ant-total 0.633 0.729 0.393 Total insects 0.593 0.487 0.526 Orthoptera 0.545 0.378 0.528 Thysanoptera 0.470 0.756 0.138 Isoptera (canopy) 0.417 0.140 0.496 Psocoptera 0.398 0.148 0.457 Coleoptera 0.312 0.458 0.127 Hymenoptera 0.302 0.446 0.129 Formicidae 0.274 0.370 0.142 Acari 0.190 -0.232 0.443 Spiders 0.186 0.307 0.050 Blattodea 0.124 -0.014 0.204 Hemiptera 0.098 0.229 -0.026 Diptera 0.038 0.404 -0.197 0.599 0.347 0.704 Bird total spp. 0.796 0.558 0.909 Above-ground carbon # Shaded areas with r = >0.500. Bold type = high indicator value. ASB SLUM Mtg Chiang Mai 11 Nov 01 Above-ground carbon and species: PFTs along a gradient of Land Use Types, Jambi [r2 = 0.814] 60 (Y=13.56 23.52X + 11.30 X**2) 1 50 AG-carbon 2 5 40 30 4 20 9 6 10 12 8 7 14 15 13 0 3 16 1.0 1.2 1.4 1.6 1.8 2.0 2.2 Species : PFTs ASB SLUM Mtg Chiang Mai 11 Nov 01 10 11 2.4 2.6 2.8 3.0 Aboveground Carbon and Species:PFTs All ASB benchmark sites Plant Spp:PFT ratio 3.00 Primary Forest 2.50 2.00 Managed Forest Fallow 1.50 Tree-based 1.00 0.50 Crop Pasture 0 50 100 150 200 250 Aboveground - C t/ha 300 350 400 Correlations between soil and vegetation variables* Variable pH-H2O pH-KCl C% N% C/N% Na Al3 H+ PFT -0.448 0.028 -0.459 0.024 0.586 0.003 0.592 0.002 0.497 0.013 0.568 0.004 0.465 0.022 0.503 0.012 Species -0.424 0.039 -0.427 0.037 0.529 0.008 0.585 0.003 0.435 0.033 0.503 0.012 0.458 0.024 0.441 0.031 Mean Height -0.358 0.086 -0.331 0.114 0.559 0.005 0.530 0.008 0.477 0.019 0.457 0.025 0.379 0.068 0.480 0.018 Basal Area -0.331 0.114 -0.260 0.219 0.482 0.017 0.572 0.003 0.383 0.064 0.375 0.071 0.400 0.053 0.402 0.052 V-Index -0.433 0.034 -0.432 0.035 0.550 0.005 0.622 0.001 0.448 0.028 0.495 0.014 0.486 0.016 0.457 0.025 Tree wt1 -0.588 0.002 -0.586 0.003 0.777 0.000 0.566 0.004 0.722 0.000 0.700 0.000 0.570 0.004 0.755 0.000 Tree wt2 -0.576 0.003 -0.568 0.004 0.753 0.000 0.557 0.005 0.697 0.000 0.668 0.000 0.558 0.005 0.735 0.000 Plot Age -0.421 0.041 -0.410 0.046 0.613 0.001 0.566 0.004 0.528 0.008 0.569 0.004 0.390 0.060 0.546 0.006 * Upper line = ‘r’ value; lower line = ‘P’ value; shaded areas with P <0.020 Tree wt1 = Quirine data; Tree wt2 = Brown data; only variables with highest correlations listed.(Source: Gillison, 2001; Hairiah and van Noordwijk, 2001) Conclusions • Specific knowledge of functional types may be essential to predict ecosystem responses under different global scenarios or where management seeks to manipulate species composition directly as in complex agroecosystems. • Hypotheses and models must be tested in a wider array of ecosystem types e.g. tropical forests. • To predict and understand changes in biodiversity and ecosystem function we need to move beyond simple causality and address multiple feedbacks. • Relationships between local, landscape and regional scales require critical attention. ASB SLUM Mtg Chiang Mai 11 Nov 01