Many ecosystems can bbe considered to be large food webs. C HAPTER 20 Trophic Structure Road Map 1. Food webs connect ll species that feed on one another in an ecosystem. 2. Generally, the connectance remains constant between different food webs, but the mean cchain length increases as the number of species in the web increases. - Problems such as imperfectly known links between species, the taxonomy of species, and the amount of prey eaten by each species can profoundly affect food web patterns. |. Ecological guilds are groups of species that feed on the same resources in the same way, and the species involved are likely to be competing for resources. Keystone species have an effect on an ecosystem out of all proportion to their biomass, 'n 1935, the British plant ecologist A. G. Tansley coined the term ecosystem, which ‘he took to include not only the biotic com- ‘munity of organisms in an area, but also the abiotic environment around that communi- ty. Ecosystem ecology concerns the move- ment of energy and materials through communities, and like the concept of a com- ‘munity, the ecosystem concept can be applied at any scale: A drop of water inhabited by protozoa is an ecosystem, and a lake and its biota are another. Most ecosystems can never really be regarded as having definite bound- aries. Even in a clearly defined pond ecosys- tem (Fig. 20.1), waterfowl may be moving in and out, The big advantage of studying ecosystem ecology is the common currency of energy or nutrients, which allows the biol- ogy of communities and populations to be compared between and within trophic levels, 20.1 Food Web Complexity Few ecosystems are so simple as to be char- acterized by a single unbranched food chain in which one plant species is eaten by one herbivore species that is in turn fed upon by one species of carnivore. Usually, ‘many herbivore species feed on the same plant species. For example, one may find tens to hundreds of insect species, as well as several vertebrate grazers, all feeding on ‘one species of tree. Also, many species of herbivores eat several different plant species, not ust a single one, Such branch- ing of food chains occurs at other trophic levels as well. For instance, frogs eat many different types of insect, and owls may eat herbivores such as field mice and also predatory organisms like snakes. It is more correct, then, to draw relationships between these plants and animals not as a simple chain, but as a more elaborate interwoven food web, as illustrated in Fig. 20.2. The first measure of the relative com= plexity of a food web can be described by a measure known as the chain length, which denotes the average number of links be- tween trophic levels. So, if there are three trophic levels—plant, herbivore, and cari- vvore—there are two links between the three species, and the chain length is two. The sec- ond measure is connectance: actual number of links connectance = tential number of links@ “Lewepeusioiy 0} foseq © ‘£ emrpeuse}4 0) ew}PEuLeRUL '5 dou oy reseq $1.60) 01 epost 106009 Jo 2sqUNW 1881 s990d8 eye)peUueyU jo JINN, (EE REOTEET e000 «| overdo equnn "te igor essay poet Bt mo Sie) sam area 2 1): ds onpnouy tr , ds srilsoxivedon (0) ‘aconivedeu vous (6 wera +) Neve sopoed f) “wayewelynyos euyjuedeuopuz (9) ‘feumuedu ‘ds eyasebayy (¢) ‘SrédoGuuAs sisojd)po1se7 (p) . {88094 se1yoUutyox0y (6) " 41 48@u) ppukou3 (2) ‘B}oOKaUedeU ScouBLUNsipy (1) ‘Key, f | (2uy ‘saxaunasey oroyg ‘silog 8 JAD) "ssAeUR gam poo} 01 JOM s2npSUIBLA Buoy wou (e) ui panuasaudas asoya a4 ‘stuanshso29 PPaurewo>-y98 umo aye aney sn oH ued JaKIg "eshejery “Guewug ye 33104 auUOWN UI 'auo}opoW Seuiuedn ued Jou (Q) (596 128099 uy) ‘ford say ven ainy aif 442464 aie sone Paid ‘960 1ydon e syuasaidas auy ype “ashe 159) Ut DiouBsowogpo saxnuaday quel y>nd 34h 1 spasur sta Jo Gam poo, aiduus ¥ (e) z-9z aang siotunsuo> Kie “tua 24g armpsuo> pue ysy ays axa uo poo sajum 959 sua ut 40 sy 19640) 4 “feu "sownsu09 Aiepudras Bavapisuo> aq ue> pue uot juerdooz pue si22sul aug uo Pap) Ys yews “saoNqioy 20 siawunsuo> Ksewd auf are 522541 10 (uoyyueydoo) sewue a1do2s0.9y "uaDnp ord Lewd ap 97 sued 21001 40 ued ado>s ony) sarreds wuaiayip e ‘4 patdn220 51 an} >yydon, Ype3 warshs0>2 puod UL gam Pood Loz aanb1g ete 2 ‘slleyd wynasen “siaonpoig ££ aimanns aydoy,338 SECTIONS. 10,000 1000 100 Number of links, 10 on Patterns in food webs focus on connectance linkage densities. Log mean chain length Figure 20.4 The length of food chain increases asthe num ber of species in the food web increases. (Modified from Martinez values, chain lengthe, and 1.00: 7s: 00. 025 Ecosystems Ecology 10 100 Number of species Figure 20.3 Food web connectance. The gray line gives the line of con- stant connectance such that the number of links in a food web goes up Lniformly with the number of species in the web, The re line gives the line ‘of best fit for data from 175 actual food webs. Note thatthe scale on both ‘axes is logarithmic, (Modified from Martinez, 1992.) Inan ecosystem of n species, the number of potential links (assuming that links cannot 0 in both directions) is _ mn = 1) NO 2 ‘This means that each species can eat any other species, but “prey species” cannot tum os 19 1s 20 around and eat their predators, which seems logical. In the pitcher plant community, there are 19 “species,” 33 actual links, and 19 x 18/2 = 171 potential links. Therefore, the con- nectance ofthis community is 33/171 = 0.19. The third measure of complexity is the number of links per species, called the link- ‘age density. For the community in the pitcher plant, the linkage density is 33/19 1,74, The three measures of chain length, connectance, and linkage density are wide- ly used to describe the complexity of food ‘webs and to compare complexity between different types of food webs. 20.2 Food Web Patterns Using data on connectance and linkage den- sity, ecologists have put forth a number of ‘generalizations about food webs. Most of these generalizations have been developed from an analysis of published works on food webs. Perhaps the most important as- sertion is that connectance remains constant ‘as the number of species in the food web in- ‘creases (Fig. 20.3). A constant level of con- nectance can be understood by imagining, an insectivorous bird that feeds in two com- munities, A and B, with A having twice the number of insect species that B does. Neo Martinez, of San Francisco State University (1992), argued that it is then reasonable to suppose that the bird would eat twice as ‘many species of insect in community A as in community B. Because this conclusion ‘would likely apply to all species in the com- ‘munity, the connectance would remain un- changed as the species richness increased. Measures of connectance from 175 real food ‘webs support this reasoning. ‘Another important pattern is that the ‘mean chain length increases as the number of species in the web increases (Fig. 20.4 Although this relationship is intuitively ob- vious, it probably has an upper limit, so that the chain length reaches a maximum; we just don’t know where this maximum is yet Finally, a long-known pattern is that top predators tend to be rather large and sparse- ly distributed, whereas herbivores are smaller and more common. This general- ization is often termed the pyramid of num- bers or, sometimes, Elton’s pyramid, in deference to British ecologist Charles Elton, who first described it in 1927. Elton’s ex- ample was that of a small pond, in which the numbers of protozoa may run into the millions and those of Daphnia and Cyclops (their predators) into the hundreds of thou-uuesKd sy "300 60 Ue UO ‘Gury sustueBi0 uo paseq ‘ssequinu jo paweskd pavon ut (@) Yor9| auydon Bussea> “ur uum sosea1D9p puod snug fed © u sapads 40 22uepungy (2) “vaquinu 430 spuwekd $"0z San6tg 68E —aumpnns >14dou OZ WALAVHD ‘seuvot og - 281 480 8g - sesonaien pk comer 3 Bp onp senin-cw.te9 en > ‘ay Se posn st sseuro1g uy asa M990 [TAS ueo sprurezAd payraau ss2jau2UON “OS0z “8p wondaoxs eae 3ou st 998 eo aM FEN smoys jana] aydouy yoee ye ‘suisTue=L0 JO sroqumu wey Joypes ‘sseur ayy ye SUP{OO] °8 001 anoge ins ssoyepaud ay pue “By ¢ TRIO} aan at UO Saxoaiquay auf [Te BY 000'0E syBiem 908 eo ay Today aIYdon Yea UT sursmue8i0 axp ySro% 0} st prureaAd s 20313 yung eu - ysiy v0 - 6a ¥e0) 0) uondaoxa juaredde snp apouosa: of Aeon 3894 aL “(AS 02 Sta) sauisezed pure sioy -epaid jo spuesnotp jo suay troddns wim ut pya ‘sraumsuo Areunad saxo pur ‘sze| lio ‘sopj20q snozoaigiay jo spuesnoy, suioddns ‘2onpoid auo ‘aan eo uy “pri -wakd snp 04 suondaoxa ye2aaas 0 4uNA UD 8UQ “(8S'0Z “BEx) YSY DOL Pue aearey apaeq 00001 Atasssod 2q Thm aarp seazayan ‘spurs340 SECTION 5 Problems with food web patterns focus on Iimperfectly known numbers and strengths of linkages. Ecosystems Ecology ‘measure: In the English Channel, the bio ‘mass of phytoplankton supports a higher biomass of zooplankton (Fig. 20.54). This is possible because the production rate of phy- toplankton is much higher than that of 200- plankton, and the small. phytoplankton standing crop (the biomass at any one point in time) processes large amounts of energy (Fig. 20.5e). The most realistic pyramid is thus the energy pyramid, which never be- ‘comes inverted. 20.3 Problems with Food Web Patterns While food web theory has undoubtedly led to an increase in the understanding of ‘ecosystems, many authors have pointed out numerous problems with the data that may invalidate the observed patterns: 1. Predation on “minor” species is fre- quently omitted from the literature. Often, authors of published works on food webs are not sufficiently knowl- ‘edgeable about organisms outside of their areas of specialty, so that some links in the web are never drawn. Gary Polis of Vanderbilt University in Ten- nessee (1991) argued that food webs in’ the real world are much more complex than is reported in the food web litera- ture. In a relatively simple desert sys- tem in the Coachella Valley, California, Polis conducted an intensive 10-year study of the local ecosystem and found 174 species of plants, 138 vertebrate species, 55 species of spiders and scorpi- cons, an estimated 2,000 to 3,000 species of insects, and an unknown number of nner Sneed, microorganisms and nematodes, al in a single food web: Polis compared his food web with others reported in the lit- ‘erature, using statistics compiled by Joel ‘Cohen and colleagues (1990; see Table 20.1). Chains. were longer, omnivory ‘was common, and linkage density was ‘much greater in the intensively studied Coachella food web. Goldwasser and Roughgarden (1997) suggested that ‘most food webs appearing in published ‘works are too poorly sampled to be use- fulin developing theory. By using com- puter simulations, the authors were able to randomly sample a fully known Caribbean food web. Food web proper- ties varied markedly with the intensity of sampling. Data on quantities of food consumed, indicating the thickness of connecting links, are usually absent from published works. Marine ecologist Robert Paine (1992) provided the first example in which interaction strengths were calcu- lated in a food web. Most of the con- nections were found to be weak, so the ‘web was essentially very simple. This objection is virtually the opposite of the problem just discussed—there may be many more links than we realize, but most of them are unimportant. Benke and Wallace (1997) also showed how misleading a normal food web was for a riverine system in Alabama, becauseit implied the equivalence of all food re- sources. An energetic analysis that looked at ingestion rates of food actu- ally showed great variation in the strengths of linkages (Fig. 20.6). 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(1) Calamomyia ateitorae (Diptera). (2) Mordelistena splendens (Coleoptera). @) Longuria taedata (Coleoptera). (4) Chilo lejadelus (Lepidoptera), (5) Thryptcus violaceus . (Diptera). Arrows indicate wherein the stem the larva lof each species is found and the drection it bores. ‘Adults, shown at extreme right, are free living. The length of each scale line be- tween the larva and adult of each species represents 0.5 min the drawing, SECTIONS Ecosystems Ecology (1967) in his studies of birds and was meant to describe a group of species that fed on the same resources and in the same way. For instance, in an insect community feeding on a plant, we may have the leaf-chewing guild, the sap-sucking guild, leaf miners, stem borers, stem gallers, root feeders, and flower feeders. There are five species that feed by boring into the stems of salt-marsh cordgrass, Spartina alternifiora (Fig. 20.7) Why should we be interested in guilds? ‘There are two main reasons. First, guilds ‘might represent arenas of the most intense competition. Attention is focused on all competing species, regardless of their taxo- nomic relationship. For example, competi- tion in a seed-eating guild may involve rodents, birds, beetles, and ants. Thus, for completeness’ sake, investigations on com- petition should involve all guild members. This forces researchers to cross taxonomic lines. Second, and perhaps more important, guilds might represent the basic building blocks of communities. Eric Pianka of the University of Texas (1988) envisioned a pe- riodic table of guilds, analogous to the pe- riodic table of elements, with entries such as insectivores, granivores, and so on. Pre- sumably, one might be able to predict inter- actions between guilds in much the same ‘way as we can predict interactions between elements in different families of the period- ictable, although ecologists haven't built up to this level of sophistication yet. ‘The main problem in guild theory is ac- curately defining the guild. How much overlap in diet does there have to be for ‘species to belong to the same guild? For the Spartina stem-boring guild illustrated in Fig. 207, there is no problem: All species feed on the inside of the stem. 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Dots represent species and areas of oak leaves on which they feed. Lines connecting dots represent species that feed in more than one area. There are many similarities between the : ‘two countries inthe area of the oak leaves where each guild feeds and in the number of species In each guild. (After Ashbourne and Putnam, 1987.) Keystone epecies are members of a community ‘that have an effect ous of proportion to their abundance. guilds did not exhibit any constancy be- tween geographic areas. For example, there are no miners in England, but many in New Guinea (Fig. 15.10). The results of Lawton’s study failed to support guild theory. S. R. Ashbourne and R. J. Putnam of Southampton University in England (1987) did the same thing as Lawton, using the in- sect herbivores feeding on red oak (Quercus rubra) and aspen (Populus tremuloides) in Canada and the United Kingdom. They used the same four basic guilds as Lawton (chewers, suckers, miners, and gallers) and included three more: species that folded one leaf in half and fed inside, species that tied two leaves together and fed inside, and species that rolled leaves up and fed inside. Their result was the opposite of Lawton that striking similarities were seen between the guild composition of the two regions ig. 20.9). The two results are typical of what has been found so far: Some studies, find similarities in guild structure between ecosystems, and others do not. As yet, there is no consensus regarding which pattern is the more frequent. 20.5 Keystone Species Within ecosystems, food webs, and even guilds, some species may have an effect out of all proportion to their abundance or bio- mass. For example, the beaver, a relatively small species, can completely alter an ecosystem by building dams and flooding anentire river valley. Al the trees in the val- ley die, and fish and wildfowl may become common in the resultant lake. This patterncontrasts with patterns in which the impor- tance of species in an ecosystem varies ran- domly or in which each species is equally important (Fig. 20.10). The removal of species like beavers obviously has severe ramifications for a majority of community members, promoting fish die-offs, loss of waterfowl, and the death of trees adapted to waterlogged soil. The term for such species, keystone species, has enjoyed an enduring popularity in the ecological liter- ature (much like the term guild) since its in- troduction by Robert Paine in 1969. It is important to distinguish a keystone species from a dominant species. A dominant species is a species that has a large effect in a community because it is very common. For example, Spartina cordgrass is a domi- nant species in a salt marsh in virtue of its large biomass and role in energy flow. Other common dominant species in different ecosystems include trees, prairie grass, corals, and giant kelp (Fig. 20.11). Several different types of keystone cate- gories have been recognized: keystone ene- mies, keystone prey, and keystone habitat modifiers. Among the most famous keystone predators are Pisaster starfish and predatory Whelks (Conchloepas) in the rocky intertidal zone (Paine 1966, 1969). Removal of these camivores leads to the nearly complete dom- inance df the substrate by one or two species cof mussels that outcompete the other con- sumer species, resulting in greatly reduced species diversity, as we saw in Chapter 15. A. rare rhinovirus that makes a wildebeest sneeze, Ve, would not qualify as a keystone species, because its impact is relatively low, buta rare distemper virus, Vo, that kills ions or wild dogs, would, Sea otters have been la- beled as keystone predators because they limit the densities of sea urchins, which, in tur, eat kelp and can destroy this dominant species in the absence of the otters. ‘Tropical ecologist John Terborgh (1986) considered palm nuts, figs, and nectar to be keystone prey because they are critical to tropical forest fruit-eating guilds, including primates, rodents, and many birds. Togeth- er, these vertebrates account for as much as three-quarters of the forest bird and animal biomass. Without the fruit trees, wholescale extinction of frugivores would occur. In the southeastern United States, go- pher tortoises have been regarded as key- stone species because their burrows provide homes for an array of mice, possums, frogs, snakes, and insects. Without tortoise bu CHAPTER 20. Trophic Structure (@) Keystone pattern a5 la (©) Random pattem (Community impertance [Eh "il (Uniform pattern 3.9 8c 87s TE ATT ‘ABCDEFGH| JKLMNOPORST. Species Figure 20.10 Community importance values (le, whether or not a species has a large impact fon community diversity, nitrogen uptake, of some other function) for species in a hypothetical ‘community, (a) based on the keystone-species ‘model, (b) based on a random patter, and (c) with al species equally important. (After Mils et al, 1993.) rows, many of these creatures would be un- able to survive in the sandhill areas where they are found. The gopher tortoises act as keystone habitat modifiers (Fig. 20.12). In fact, both beavers and gopher tortoises have been called ecosystem engineers because they modify the habitat and cause ecologi- cal changes (Lawton and Jones, 1995), and this term seems to have caught on more than keystone habitat modifier. African ele- phants may also act as ecosystem engineers, through their browsing activity. Elephants destroy small trees and shrubs when brows- ing and can change woodland habitats into grasslands. Ungulates that graze the grass- es are favored by the elephant's activities More examples of ecosystem engineers and their effects on ecosystems are given in Table 202, Are keystone species present in most communities? It is too early to tell. Indeed, few studies have analyzed the community importance, or interaction strength, of species. However, Paine (1988) suggests that humans, introduced pests, and diseases 345346 SECTIONS Ecosystems Ecology W- Whe Disaster Vp - Vie, Distemper Vp - Virus, Rhinovirus Bias Ss 0: Otter K- Kelp Coral Grass Tree Proportional biomass of species Figure 20.11 Keystone species are those species whose effects are out of all pro: Portion to their biomass. Dominants are species that dominate community bio ‘mass and whose impacts on an ecosystem are large, but not disproportionate to their biomass, (Atter Power and Mils, 1995.) may commonly act as ke tone species. The of constant connectance in stich webs. How keystone species ike the beaver and the go-___ tial pitfalls of imperfectly known web To many species tha live alongside it tain parts ofthe webs, suchas guilds Key Figure 20.12 Gopher tor’ "Food web theory has leo some alr stone spcte offer gest prone te fora tose burows in US. seu able insight, suchas that into the pattem points or conserraton ain waite whch use em nginetn Effect en Sevag aan Soll enyankans Inctease and eduction of andar utes bemeet Atredycology Cora eet Physica structure alerajon of ove ates i Proll sirvcture oper trots Biel Woodpecters Nest cavity Terie mitnds Homes fo other antvopods Vertebrate feces Homes for dung beetlesSummary ai CHAPTER 20 Trophic Structure 347 Ecosystem ecology concerns the flow of energy and nutrients through communities. It offers the use of a common currency—energy or nutrients—by which to compare different ecosystems, 1. Many food webs can be compared by simple properties such as connectance (degree of connectivity) and linkage density (numberof links per species). 2. The analysis of food webs has revealed some common patterns. Two are that, as the number of species in a food web increases, chain length tends to increase, but connectance remains constant. Another is that there is a pyramid of numbers, with fewer species on higher trophic levels than on lower trophic levels. 3. Because food webs are usually imperfectly known, the generalizations that have resulted from analyzing them may be incorrect. The inadvertent omission of minor or mobile species, lack of knowledge about the strength, importance, or beneficial effects of connecting links between species, and the taxonomic lumping of species into groups or guilds can all obscure true food web patterns. 4. Itis possible to split up trophic levels into units called guilds For example, among herbivores, we can recognize the leaf- chewing guild, the stem-boring guild, and the root-feeding guild, Guilds area valuable analytical tool because they focus attention ‘on groups of species most likely to be competing for resources. 5. Keystone species have a large influence on ecosystems out of all proportion to their abundance. A common keystone species is the beaver, whose activities can flood entire valleys and completely change the composition of the species that live there. Food webs offer a potential shortcut in understanding how communities work, However, to yield reliable insights into the way a community functions, food webs must be thoroughly known. Discussion Questions 1. How might food web properties change if we included all the decomposers in food webs? Much energy passes through ecomposers. 2. Do you think food webs would be different in complexity between two-dimensional habitats, such as grasslands, versus three-dimensional habitats, like forests? 3. Would you expect differences in chain lengths between disturbed and undisturbed areas? How would disturbance affect conneclance of linkage density? 4. Would fertilizing an ecosystem increase the length of a food chain, and if so, why? 5. Do you expect the position of a species in a food web to change as it ages and increases in size? Ifso, does this affect how we construct food webs?Five ways of measuring primary production are by calorimetry, harvesting, carbon dioxide uptake, oxygen output, and chlorophyll concentration 348 C H A=P_T 3E=R Energy Flow Road Map 21 1. The energy content of plants can be measured by calorimetry, harvesting, CO; produc- tion, O2 production, or chlorophyll concentration, 2. The production of plants is limited by temperature, moisture, and nutrients such as nitro- gen and phosphorus. 3. Primary production is highest in the tropics and decreases toward the poles. 4. The efficiency of the transfer of energy from plants to animals can be measured by three indices: production efficiency, consumption efficiency, and trophic-level transfer efficiency. 5. The production of animals is limited mainly by the production of plants. ‘Pietsrsomer naomi. ca Fis cin be woe asa vest Ouamon denominator and popuitons canbe fe ded tcaoricequtents nthe seach or jection ingodogy.onexeyensep proach may provide nights no how be Dope tee function by lsely examining theenery content diferent tepclevels Energy sntent generally meer wing Gy brass ody weight because he bulk conta weft ts soora DY ea ‘more meaninghl euro elo cor, tent Of the dy weight 95% i made up of carta, one at paste ey fow ineaeysiena bin many wae oe 21.1 Measuring Production ‘The bulk of the Earth's living mantle, 99.9% by weight, is green plants; only a small frac- tion of life is animal. Therefore, when we measure the energy of an ecosystem, we are interested primarily in plants. Because plants represent the first, or primary, trophic level, Wwe measure plant production as gross pri- mary production, which is equivalent to the energy fixed in photosynthesis. Net primary production is gross primary production ‘minus the energy the plant loses through res- piration. Net primary production is the amount of biomass accrued by plants, meas- ured as the weight of living plant material, including leaves, stems, and roots. In order to examine ecosystems as en- ergy transformers, we must have some way of determining the energy content of plants. This can be done in many ways, and we will examine five of the most common, In calorimetry, a sample of dry material is burned in a small chamber, where it oxidizes completely to carbon dioxide and water. High-pressure oxygen is fed into the cham- ber to ensure complete combustion. A water jacket surrounds the chamber, and the rise in temperature of the water provides a measure of the heat generated by combustion of the sample. The greater the energy ina sample, the more heat will be generated. In harvesting, samples of plant biomass are obtained at the beginning of a growing season, after which they are dried andweighed. At the end of the season, another sample is obtained and then dried and weighed. The difference of the two weights represents growth in plants in the area stud- ied and provides a measure of pro ina known unit of time In the CO; uptake method, plants are grown inside sealed chambers for short pe- riods. Because we know that air contains 0.03% CO, any decrease in CO; inside the chambers is the result of CO; uptake by plants during photosynthesis. The higher the CO; uptake, the greater is the produc tion of plant biomass. The exact plant bio- mass required to take up a given volume of COIs then calculated, ‘We can estimate the production of many aquatic plants by their O; output instead of CO, intake. For example, to measure phyto- plankton production, samples of phyto plankton are suspended in two bottles in a lake or an ocean. One of the bottles is dark in color, so photosynthesis is not possible and oxygen is used during respiration, The other bottleis light, and photosynthesis occurs, giv- ing off oxygen beyond that used in respira- tion. The difference in oxygen concentration between the light and dark bottles yields the total oxygen produced by photosynthesis. - = (__ Most plant biomass Seal ae x Most eneray is ost. inphotosyntnesis Figure 21.1 Energy-tlow diagram for a Georgia sal {Clos and ros in place (CHAPTER 21 Finally, production can be estimated by measuring chlorophyll concentration. We know how much carbon is assimilated per gram of chlorophyll. By taking a sample of the plant under question and measuring its chloro- phyll, we can calculate its carbon production. More modern methods take advantage of the reflectance of plants to measure the chloro- phyll concentration. Thus, orbiting satellites can identify reflectance colors of different habitats and thus estimate their production. Using these techniques, we begin to an- alyze the amount of energy contained in dif- ferent ecosystem components, such as plants, herbivores, and carnivores. To analyze a.com plete ecosystem in this way, calculating the calorific content of all the component species, isa monumental task. However, some ecolo- gists have concentrated on analyzing rela: tively simple systems such as salt marshes, in which there are few species. The patterns developed from such systems should be valid for other communities as well Among the pioneers of this type of work was John Teal, of the University of Georgia, who examined energy flow in a Georgia salt marsh (Fig. 21.1). In the marsh, ‘most of the incident energy from the sun went to two types of plants: Spartina cord- 7 {( Uitie plant biomass) eaten by herbivores J Much energyis also.) Tost in espiration it marsh. Units are klocalores per square meter per year. Thickness of flows are not exactly to scale. (Redraw from Tea. 1962.) The pho: tos shown ab ve are a Georgia salt marsh and a fiddler crab. |. and nematodes Energy Flow ‘About 1036 of the bacteria are eaten by crabs t Not 7 export 349350 SECTIONS Primary production is limited mainly by light, water, nutrients, and ‘temperature. Ecosystems Ecology ‘grass and marine algae. The cordgrass plants ‘were rooted in the ground, whereas the algae floated on the surface of the water or lived on the mud or on the Spartina leaves at low tide. The cordgrass and algae are the producers in the system. Most of the plant energy is used in the energetically costly process of photo- synthesis. The remainder is lost mostly in res- piration. The energy that is accumulated in plant biomass is small: 34,580 keal m-2 yr-1in Spartina and 1,800 keal m-? yr~"in the algae, totaling 36,380 kcal m-? yr ~!, or about 6% of the incident solar radiation. The herbivores take very litle ofthis plant production, eating conly 305 kcal m-? yr “tof the Spartina (0.8%) and none of the algae. The remaining 99.2% of the plant biomass dies in place and rots on the muddy ground, to be consumed by bac- teria, the major decomposers in the system. In tum, the bacteria are eaten by crabs and ne- matodes, which sift through the mud. Very little of this dead material is removed from the system (exported) by the tide Inthe salt marsh, the producers are the ‘most important consumers; in other words, most primary production (98.6%) is used in plant respiration and photosynthesis. The bacteria are next in importance: As decom- posers, they degrade about 20% of the ener- sy that plants use. As with many ecosystems, Production (g/m) 200 400 600 ‘most plant biomass goes to the decomposers. Animal consumers are a poor third in im- portance, degrading only about 3.8% of the energy the bacteria use. 21.2 Limits to Primary Production What limits primary production? In terres- trial systems, water is a major determinant, and production shows an almost linear in- crease with annual precipitation, at least in arid regions (Fig, 21.2). However, tempera- ture is also important. Temperature affects production primarily by slowing or accel- erating plant metabolic rates. For example, one study indicated a linear relationship be- tween total net primary production and ‘mean annual temperature, as measured on an elevational gradient in Hawaii: For each 1°C increase in mean annual temperature, total net primary production increased by 54 gm~2 yr-! (Raich et al., 1997). Michael Rosenzweig of the University of Arizona (1968) noted that the combined effects of temperature and moisture (ie., the actual evapotranspiration rate) could predict the aboveground production with good accu- racy in North America (Fig. 21.3). The evap- Precipitation (mm) Figure 21.2 Relationship between mean annual precipitation and mean aboveground net pri mary production for 100 areas across the central grassland region of the Great Plains ofthe United States. (Redrawn fram Sola et a, 1988.)CHAPTER 21. Energy Flow 351 as opie ost Dckdoue ret Fi orm tt i oe ‘Oak-hickory forest z g ‘esi NY / Sa i As «i Ba y 20 hs ; 2a eSCSC Log of evapotranspiration Figure 21.3 Plant production is positively correlated with evapotranspiration Warm, humid environments are ideal for plant growth. Dots represent diferent ecosystems, some of which are indicated. (Modified from Rosenzweig, 1969.) men chiemetese ube oa iva Gapmaniocioe ceeentiiae SE gon creates teropeaten toe Be Freniiniciarecicrren ates Se repeaiiaranitteampneccheciow’ Be wiinivelow cropereeena tors | reuters sla ete gh te patra bay when hed oars an [> Tete wetter opt ionamitien Highemmomttiea Pyaar ecetearpaese ac Sion en Saviotoa el Tn adclicn we neoclsion th as restucsndcoicure ean Fsclonaffncarielloeined ote jeg ofthe gootiog seeobtinorne days available for forests to grow, the more wood they can put on. For this reason, southern Appalachian forests of Tennessee and North Carolina are more productive than their northern counterparts in Wis- consin and New England. The length of the growing season helps explain why tropical wet forests, with a very long growing sea- son, are so productive and why deciduous trees, predominant in northern realms, are not. However, conifers at northern latitudes effectively extend the short growing season at high latitudes by retaining their leaves (needles) for long durations. Nutrient deficiency, particularly of ni- trogen and phosphorus, can limit primary production also, as agricultural practi tioners know only too well. Fertilizers are352 SECTIONS. Primary production in aquatic systems is limited the availablity of 300 200 100 Ecosystems Ecology commonly used to boost the production of annual crops. Stewart Cargill and Rob Jef- fries (1984), of the University of Toronto, showed how both nitrogen and phospho- rus were limiting to salt-marsh sedges and ‘grasses in subarctic conditions in Hudson Bay, Canada (Fig, 21.4). Of the nutrients alone, nitrogen was the most limiting: The addition of phosphorus failed to increase production. However, once nitrogen was added and was no longer limiting to the system, phosphorus became the limiting factor, and the addition of phosphorus and nitrogen together increased production the nost. This result supports Justus Leibig’s 1842) law of the minimum, discussed in Chapter 7, which says that the abundance of species is controlled by the most limiting factor—in this case, nitrogen. The most lim- iting factor can change, as in the Hudson Bay marsh: When sufficient nitrogen is available, phosphorus becomes limiting. ‘Once phosphorus becomes abundant, pro- dluction is still ikely limited by some other Nitrogen and phosphorus ‘Control Phosphorus Nitrogen only ‘only Treatment Figure 21.4 Net aboveground primary produc tion of a salt-marsh sedge, Carex subspathacea, in Canadian salt marshes in response to the addition of nutrients, Nitrogen is more limiting than phos. phorus, but ance nitrogen becomes available, phosphorus then becomes limiting, (From dota in Corgi ond jetties, 1984.) nutrient. Apparently, there is a sequence of limiting factors in ecosystems. Aquatic systems. (Of the factors limiting primary production in aquatic ecosystems, among the most im portant are light and nutrients, Light is par ticularly likely to bein short supply, because water readily absorbs light. At one-meter depth, half the infrared energy has been ab sorbed, By 20 meters, only 5 to 10 percent of the radiation is let. Too much light can also inhibit the growth of green plants, by ‘overheating them. Such a phenomenon can be found in tropical and subtropical surface waters throughout the year; maximum pri- mary production occurs several meters be- neath the surface. The most important nutrients affecting primary production in aquatic systems are again nitrogen and phosphorus. Important only locally in terrestrial systems, both often limit production over large areas of the oceans, where they occur in very low con centrations. Few nutrients are tied up in the standing crop in aquatic systems, in contrast to terrestrial systems: Soil contains about 0.5% nitrogen, whereas seawater contains only 0,00005% nitrogen. Enrichment of the sea by the addition of nitrogen and phos- phorus can result in substantial algal blooms. Such enrichment occurs naturally in areas of upwellings, where cold, nutri- ent-rich, deep water is brought to the sur- face by strong currents, resulting in highly productive ecosystems and plentiful fish Such upwelling occurs in the Antarctic and along the coasts of Peru and California, and these are rich, productive waters. Phosphorus is particularly important in limiting. production in freshwater lakes Some lakes in North America become pol Iuted by runoff of rainwater enriched with phosphorus from fertilizers or from sewage. This input into the lakes causes h of algae, which clog the lake and increase its turbidity, a process termed eutrophication, One of the champions of the study of eu trophication is David Schindler (1974, 1977), who experimentally added phosphorus to lakes in northwestern Ontario. Not only does phytoplankton biomass increase with increased phosphorus (Fig. 21.5), but also, the composition of the algal community changes from green algae to blue-green algae. The blue-green algae can produce some secondary chemicals, so zooplankton50 (maim) 30: Mean annual chiorophy prefer the green algae. Asa result, the phos- phorus-laden lakes become clogged with a scum of blue-green algae that is reviled by fish and humans alike. The degree of eutrophication of large bodies of water like the Great Lakes shows a striking resemblance to maps of human population density along the shoreline. In- dustrial and urban wastewaters, especially those containing phosphorus, cause mas- sive eutrophication. Even rural areas with high concentrations of livestock can cause eutrophication of water bodies as agricul- Relative uptake of carbon (ameasure of productiviy: CHAPTER 21 70 20 30 0 Mean annual total phosphorus (mg/m) tural wastes seep into the water. The reme- dy, of course, is to divert wastewater away from lakes, or else to treat the wastewater, removing phosphorus and nitrogen before they discharge into lakes, In marine systems, nitrogen and phos- phorus also limit production, and iron is known to be limiting as well. Large-scale ‘open-ocean fertilization experiments in the Sargasso Sea in the early 1960s showed that the addition of iron to surface water boost- ed production, as revealed by an increase in carbon uptake (Fig. 21.6). _ (Cen limits phytoplankton productivity, lL Metals NePs NeP N+P+ N+P+ allmetais only allmetals Iron ‘except Treatment Energy Flow 353 Figure 21.5 Increase in lake primary production, ‘measured as chlorophyll concentrations, with phos- phorus concentration, More ‘algae grow as phosphorus levels increase. (Ater Schindler, 1977.) Figure 21.6 Resi of ex- periments in wich bottled Seawater fom the Sargasso Sea, south of Bermuda, was enriched wit various nut ents, Production in exped- ‘mental bots was compared relative to that in controls to hich no nutrients were added. ron was the limiting factor. From data in Menzel ‘nd Ryther, 1961.)a 354 SECTIONS The efficiency of primary production can be calculated by comparing ‘the energy produced by Photosynthesis with the ‘energy available in incident sunlight. Se as Deciduous trees Ecosystems Ecology 21.3 Patterns Production Efficiency of primary production. Armed with the knowledge that precipitation and temperature are limiting factors, we can ‘make some predictions about the efficiency of primary production. For example, we could ‘expect deserts to be not very efficient at con- verting huge amounts of incident energy from sunlight into plant biomass, because water is scarce. In fact, one can determine the efficien- cy of utilization of sunlight from the ratio Primary efficiency of gross _ primary production ~ energy fixed by gross prim: Phytoplankton communities have very low efficiencies—usually, less than 0.5%—be- cause they are severely limited by the avail- ability of nutrients (Fig. 21.7). The highest efficiencies occur in coniferous forests, be- cause their numerous needles present a large surface area for photosynthesis. A higher efficiency generally leads to a high annual primary production. Thus, herba- ceous and forest communities are efficient and highly productive, whereas desert and tundra ecosystems are not. So far, we have looked mainly at pat- terns in gross primary production, but the patterns for net primary production are similar. Perhaps 50 to 70% of the energy fixed by photosynthesis is lost in respira- tion, so usually less than 1% of the sun’s energy is actually converted into net pi mary production. In the temperate zone, this 1% is equivalent to 300 to 600 calories of primary production per square meter of land per year. Efficiency of gross primary production 0 0s 10 Phytoplankton 182028 ao wee a f ] _Wianiyeae) ees ~acks to your suggestions? 3. What limits the efficiency of secondary production? Are native herbivore-rangeland systems such as bison-prairie likely tobe more efficient than exotic systems like cattle-grassland?