Flow-Driven Variation in Intertidal Community ...

Flow-Driven Variation in Intertidal Community Structure in a Maine Estuary George H. Leonard; Jonathan M. Levine; Paul R. Schmidt; Mark D. Bertness Ecology, Vol. 79, No. 4. (Jun., 1998), pp. 1395-1411. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199806%2979%3A4%3C1395%3AFVIICS%3E2.0.CO%3B2-Y Ecology is currently published by Ecological Society of America.

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Eci,lojiy, 79(4). 1998. pp. 1305 141 1 0 1998 b y the Ecolog~calS ociety of Arnerlc,~

FLOW-DRIVEN VARIATION IN INTERTIDAL COMMUNITY STRUCTURE IN A MAINE ESTUARY GEORGEH. LEONARD, J O N A T I ~ AM. N LEVINE,' PAULR. SCHMIDT,A N D MARKD. BERTNESS Brown Unii,ri-sity, Departmerzt oj'Ecology and Evoluriorznry Biology, Providence, Rhodr I,slarzd 02912 USA

Abstrcict. Understanding the factors that determine community structure remains one of the most important issues in ecology. In this paper, we examine the role of flow velocities in governing community structure in marine intertidal communities. We broaden the traditional definition of "bottom-up" forces to include the delivery of nutrients, food, and larval resources to habitats and then test the hypothesis that, by controlling these fluxes as well as mediating predator effects, flow velocities leave strong bottom-up signatures on shoreline communities. We examined this hypothesis by quantifying community structure and dynamics at high and low flow sites in a tidal estuary in Maine. High flow sites were characterized by dense barnacle and mussel cover, while low flow sites had considerable bare space. High flow sites also had greater grazer and predator densities than low flow sites. Recruitment of all common shoreline organisms with planktonic larvae was greater at high flow sites, in direct proportion to the increased fluid flux there. Flow effects on the growth of the herbivorous and carnivorous components of the food web were less predictable. High flows increased the growth of barnacles, but not mussels, and increased the growth of the carnivorous gastropods that preyed on them. In contrast, high flows decreased the accumulation of benthic diatoms, but this was unrelated to the growth rates of herbivorous gastropods. High flow sites were universally characterized by low predation intensity and per capita predation rates on all three prey species. Our results show that the strengLhof top-down and bottom-up forces varies with flow rate in this estuary. Consumer stress (top-down) models accurately explain patterns we saw at low flow sites, but bottom-up processes predicted from nutrient/productivity models dominate at high flow sites. High consumer pressure is the dominant structuring force at low flow sites, whereas at high flow sites predators are inhibited, and individual recruitment and growth rates become the dominant structuring forces. We suggest that hydrodynamics may commonly decouple predation and resource processes in many aquatic systems when the physical process responsible for variation in top-down forces also acts as a strong bottom-up force. Key words: bottorn-up force.^: estuary: i~ydrodyrzczmicrefcrgr; intertidal community .structure; la,vat recruitn7ent; Maine; predcrtiorz; tidal currer?ts; top-down forces.

INTRODUCTION Understanding the factors that determine community structure is one of the most important issues in ecology. Although many factors have been emphasized (e.g., physical stress [Davidson and Andrewartha 19481, predation [Paine 19661, competition [Connell 19611, recruitment [Roughgarden et al. 19881, and primary productivity [Slobodkin 19601) their combined influences remain poorly understood. Models of community struc- ture fall into two broad categories. Nutrient/productiv- ity models (Hairston et al. 1960, Fretwell 1977, Oks- anen et al. 1981) and environmental stress models (Menge and Sutherland 1976, 1987) both seek to ex- plain similar patterns but do so from different view- points. Nutrient/productivity models predict alternating Manuscript receiied 17 July 1996; revised 16 May 1997: accepted 20 May 1997. I Present address: Department of Integrative Biology, University of California, Berkeley. California 94720 USA.

control by resources or consumers depending on the level of in situ productivity and the number of trophic levels in the food web. In contrast, environmental stress models predict control by competition or consumers depending on the level of stress and whether prey (prey stress models) or predators (consumer stress models)

are more strongly affected by this stress.

Nutrient/productivity models, originally developed

in terrestrial systems (Hairston et al. 1960), have been applied most successfully in aquatic habitats. Manipulations in streams support the predictions of these simple models (Wootton and Power 1993) and those in lakes suggest that bottom-up and top-down processes are often closely interrelated (Bergquist and Carpenter 1986, Carpenter et al. 1987). Strong predation by fishes in these systems can alter local productivity by changing the relative abundance of lower trophic levels and accelerating nutrient regeneration rates (Leibold 1989, Vanni and Findlay 1990, Vanni and Layne 1997, Vanni et al. 1997). Because these are relatively closed sys-

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GEORGE H. LEONARD ET AL.

tems, primary productivity is largely intrinsically controlled and results in a tight coupling between bottomup and top-down forces through biological feedbacks. Consumer stress models, in contrast, have been best studied and applied in marine systems, especially rocky intertidal habitats (Menge 1978a, b ) . Gradients of physical stress often affect predator mobility and per capita feeding rates and shift the primary factor governing communities from predation to competition (Menge and Sutherland 1987). The interactions between environmental stress and primary productivity in governing community structure are generally unclear (Menge and Olson 1990, Persson et al. 1996). For example, Wootton et al. (1996) found that flooding disturbance in streams, instead of shortening food chains, actually lengthens them by differentially affecting predator-resistant and predator-susceptible prey. We suggest that predictions of both nutrientlproductivity and consumer stress models may apply in systems where the physical process governing variation in resources also act as a stress (sensu Menge and Sutherland 1987) for consumers. Specifically, low productivity habitats whose consumers are also subjected to low physical stress may be structured by predation. In contrast, high productivity habitats whose consumers also experience high physical stress may be governed by strong bottom-up processes. This type of linkage between these models has not been considered previously because the physical processes operating to set nutrients in many systems do not directly affect top predators. For example, in terrestrial systems, nutrient input is largely governed by small-scale effects intimately associated with litterfall and decomposition processes (Wiegert and Owen 197 1, Hairston and Hairston 1993). In lakes, primary productivity potential is largely determined by seasonal weather conditions and local runoff (Carpenter and Kitchell 1987, Persson et al. 1992). In rivers, productivity potential is determined by flooding events that set the stage upon which fish predators then act (Power 1990). In none of these cases do these processes seem to influence consumer behavior directly. Marine systems may provide a striking contrast to these examples and provide insight into the relationship between nutrientlproductivity and consumer stress models. Until recently (Menge 1992, Menge et al. 1994, 1996) there has been little work on the interplay between resources and consumers in marine systems. This may be due to the large spatial scale over which variation in primary production operates in the sea as well as the success of consumer stress models in explaining many patterns on open coast intertidal habitats (Menge and Sutherland 1987). Marine systems, however, may be ideal to examine the relative role of bottom-up and top-down effects because of the way physical factors can drive both these processes. Many waveprotected marine systems, such as estuaries and nearshore habitats, are subjected to variable hydrodynamics

Ecology, Vol. 79, No. 4

driven largely by tides and currents. These processes can transport larvae (Genin et al. 1986, Roughgarden et al. 1988, Bertness et al. 1991, Pawlik et al. 1991), suspended food particles (Sebens 1984, Grizzle and Morin 1989, Sanford et al. 1994), and nutrients and gases (Wheeler 1980, Gerard 1982, Koehl and Alberte 1988, Carpenter et al. 1991, Patterson et al. 1991) to benthic habitats. These hydrodynamic conditions can also influence predator foraging efficiency (Kitching et al. 1959, Menge 1978a, b , Burrows and Hughes 1989) and affect top-down control. This suggests that the relative contribution of bottom-up and top-down forces in some marine systems may be related to local hydrodynamic conditions and their role as a "stress" for predators and prey. We propose to broaden the traditional definition of "bottom-up" forces for these aquatic habitats to more closely match these effects. In the strict sense, bottomup control concerns only the direct effect of nutrients on primary producers and the resulting indirect effects higher in the food web. This definition is particularly useful when there is intrinsic control of primary production in relatively closed systems such as terrestrial habitats and some lakes. In benthic marine habitats where tides and currents are common, this strict definition of intrinsic primary productivity may not apply. Productivity of these communities may be exfrinsicnlly controlled (sensu Polis and Hurd 1996) via the input of nutrients to benthic plants (Carpenter et al. 1991). Fluid fluxes in aquatic habitats also permit both new individuals (i.e., larvae) and suspended food particles to be delivered to benthic habitats via physical transport processes. The different physical characteristics of water and air (Denny 1993) and the way physical transport processes operate in these two media may represent fundamental differences in the way bottom-up forces operate in terrestrial vs. aquatic systems. The delivery of nutrients, which drives strict "bottom-up" control, is thus intimately linked to the delivery of new individuals to the food web and resources to benthic suspension feeders that often make up the base of these food webs (Paine 1966). This suggests that a broader definition, which incorporates multiple inputs to the food web, is more appropriate in these extrinsically controlled systems. In this paper, we evaluate the relative contribution of these broadly defined bottom-up processes with traditional top-down forces to community structure in an estuary in Maine, USA. This estuary is subject to highly predictable differences in hydrodynamics (tidal currents) that act at both the top and bottom of the food web. Although there has been great success in quantifying how small-scale variation in flow rate can influence recruitment rates and suspension feeding in laboratory flumes (e.g., Butman et al. 1988, Mullineaux and Butman 1991) and the mechanics of some suspension feeders are now well understood (e.g., Patterson 1991, Lesser et al. 1995), we know considerably less

FLOW-DRIVEN COMMUNITY STRUCTURE

itIig!> Flow)

7 1

1

A

Plumrnes Point utes (Low Flow) (I-Iigh Flow)

ilmmer Point

FIG. 1. Location of the six study sites along the Datnariscotta River, a tidally influenced estuary in eastern Maine. The Upper Narrows Region and Lower Narrows Region both experience elevated near-bed flow rates because of constrictions in the estuary. High flow study sites were Upper Karrows. Lower Narrows, and Hodgson Island. Low flow study sites were Upper Nart-ows, Plutntner Point Outer. and Plulnlner Point Inner.

about the food-web consequences of variable flow regimes. Ours is the first study to examine the influence of local hydrodynamics on both the herbivorous and carnivorous components of an entire food web. We examine the general hypothesis that flow regimes can play a powerful role in structuring shoreline communities. Specifically, we hypothesize that sites subjected to high fluid flow should be structured largely by bottom-up forces operating through increased delivery of larvae, suspended food, and nutrients to the benthic community. These sites arc predicted to be characterized by higher recruitment and growth rates of both benthic plants and suspension feeding invertebrates but also by low consumer pressure. In contrast, we hypothesize that low flow habitats, where currentdriven delivery rates are lower, should be structured by top-down processes because of greater predator mobility. METHOUS Study sires a n d experimental design We examined the influence of flow on shoreline community structure at six sites in the Damariscotta River, a 23 km long, tidal estuary in Maine. Freshwater input to this system is very low (McAlice 1977) and water column stability is governed largely by temperature (McAlice 1979). Wave heights are generally I 1 0 cmls and fell to 0 cmls for only a few minutes at slack tide. Maximum velocities at low flow sites, in comparison, were lower by an order of magnitude. Differences in chlorophyll a concentration among sites were negligible although higher temporal variation was apparent at all low flow sites (Fig. 3). Because of the large variation in current velocity but minimal variation in phytoplankton concentration. the flux of phytoplankton ([concentration] X [flow speed]) was much greater at high flow than low flow sites (Fig. 3 1.

30k

20

10

-i

Ecology, Vol. 79, No. 4

0

Flow High

Flow Low

Site Type FIG. 2 . Dissolution rates of calcium sulfate cylinders at high and low flow sites. Chalk blocks dissolve at a rate proportional to flow rate. Dissolution is presented as percentage loss in mass over a 5-d period. Data are ineans r 1 sb.

block erosion showed that high flow sites experienced three times greater bulk fluid transport than low flow sites (Fig. 2; F,,, = 61.97, P < 0.001). Chalk block erosion among sites was also highly correlated with both maximum free-stream velocity (range = 10-135 cm/s; r = 0.989, P I.. and S. R. Carpenter. 1986. Lirnnrlic Ilerb~vory:effects on phytoplankton populations and prirn;iry production. Ecology 67: 135 1-1 360. Bcrtness, M. D., S. D. Gaines. D. Berrnudez, and E. Sanford. 1991. Extreme spatial \ariation in the growth and reproductivc output of the acorn harriacle Se~?~ibnlunils bolciiloides. Marine Ecology Progress Series 75:91-100. Rertness, M. D., S. D. Gaines. E. G. Stephens. and P. 0 . Yund. 1992. Cor~lponentsof recruitment in populations of the acorn barnacle Srnlibalc~rr~is bcrlunoidc.$ (Linnaeus). Journal of Experi~nentalhlarinc Biology and Ecology 156: 199-215. Bcrtness, M. D., S. D. Gaines. and R. A. Wahle. 1996. Winddriven settlement patterns in the acorn barnacle, Seiizibillnrzirs btrlnrroicic..s. Marine Ecology Progress Scries 137: 103-1 10. Burrows. M. T.. and R. N. Hughes. 1989. Natural foraging of the dogxvhelk. N i ~ c t ~ l l at i p i l l ~(I-innaeus): ~~ the weather and whether to feed. Journal of Molluscan Studies 55:285295. Bustamantc. R. H., G. M. Branch. S. Eekhout, B. Robertson. P. Zouiendyk, h1. Schleqer. A. Dye. N. Hanekon. D. Keats. hf. Jurd, and C. McQuaid. 1995. Gradients of intertidal primary productivity around the coast of South Africa ant1 their relationships with consumer biomass. Oecologia 102: 189-20 1 . Hntman, C. A., &I.Frcchette, W. I< American Naturalist 118:240-26 1. Osenberg, C. W.. and G . G . Mittelbach. 1996. The relative importance of resource limitation and predator limitation in food chains. Pages 134-148 irz G . A. Polis and K. 0 .

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You have printed the following article: Flow-Driven Variation in Intertidal Community Structure in a Maine Estuary George H. Leonard; Jonathan M. Levine; Paul R. Schmidt; Mark D. Bertness Ecology, Vol. 79, No. 4. (Jun., 1998), pp. 1395-1411. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199806%2979%3A4%3C1395%3AFVIICS%3E2.0.CO%3B2-Y

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The Effect of Long-Term Alteration of in Situ Currents on the Growth of Mercenaria mercenaria in the Northern Gulf of Mexico Michael L. Judge; Loren D. Coen; K. L. Heck, Jr. Limnology and Oceanography, Vol. 37, No. 7. (Nov., 1992), pp. 1550-1559. Stable URL: http://links.jstor.org/sici?sici=0024-3590%28199211%2937%3A7%3C1550%3ATEOLAO%3E2.0.CO%3B2-7

The Ecology of Lough Ine: VIII. Mussels and Their Predators J. A. Kitching; J. F. Sloane; F. J. Ebling The Journal of Animal Ecology, Vol. 28, No. 2. (Nov., 1959), pp. 331-341. Stable URL: http://links.jstor.org/sici?sici=0021-8790%28195911%2928%3A2%3C331%3ATEOLIV%3E2.0.CO%3B2-B

Resource Edibility and the Effects of Predators and Productivity on the Outcome of Trophic Interactions Matthew A. Leibold The American Naturalist, Vol. 134, No. 6. (Dec., 1989), pp. 922-949. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28198912%29134%3A6%3C922%3AREATEO%3E2.0.CO%3B2-V

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Effects of Flow and Seston Availability on Scope for Growth of Benthic Suspension-Feeding Invertebrates from the Gulf of Maine Michael P. Lesser; Jon D. Witman; Kenneth P. Sebens Biological Bulletin, Vol. 187, No. 3. (Dec., 1994), pp. 319-335. Stable URL: http://links.jstor.org/sici?sici=0006-3185%28199412%29187%3A3%3C319%3AEOFASA%3E2.0.CO%3B2-5

Responses of a Rocky Shore Gastropod to the Effluents of Predatory and Non-Predatory Crabs: Avoidance and Attraction Peter B. Marko; A. Richard Palmer Biological Bulletin, Vol. 181, No. 3. (Dec., 1991), pp. 363-370. Stable URL: http://links.jstor.org/sici?sici=0006-3185%28199112%29181%3A3%3C363%3AROARSG%3E2.0.CO%3B2-2

Organization of the New England Rocky Intertidal Community: Role of Predation, Competition, and Environmental Heterogeneity Bruce A. Menge Ecological Monographs, Vol. 46, No. 4. (Autumn, 1976), pp. 355-393. Stable URL: http://links.jstor.org/sici?sici=0012-9615%28197623%2946%3A4%3C355%3AOOTNER%3E2.0.CO%3B2-R

Community Regulation: Under What Conditions Are Bottom-Up Factors Important on Rocky Shores? Bruce A. Menge Ecology, Vol. 73, No. 3. (Jun., 1992), pp. 755-765. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199206%2973%3A3%3C755%3ACRUWCA%3E2.0.CO%3B2-Y

The Keystone Species Concept: Variation in Interaction Strength in a Rocky Intertidal Habitat Bruce A. Menge; Eric L. Berlow; Carol A. Blanchette; Sergio A. Navarrete; Sylvia B. Yamada Ecological Monographs, Vol. 64, No. 3. (Aug., 1994), pp. 249-286. Stable URL: http://links.jstor.org/sici?sici=0012-9615%28199408%2964%3A3%3C249%3ATKSCVI%3E2.0.CO%3B2-T

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Species Diversity Gradients: Synthesis of the Roles of Predation, Competition, and Temporal Heterogeneity Bruce A. Menge; John P. Sutherland The American Naturalist, Vol. 110, No. 973. (May - Jun., 1976), pp. 351-369. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28197605%2F06%29110%3A973%3C351%3ASDGSOT%3E2.0.CO%3B2-2

Community Regulation: Variation in Disturbance, Competition, and Predation in Relation to Environmental Stress and Recruitment Bruce A. Menge; John P. Sutherland The American Naturalist, Vol. 130, No. 5. (Nov., 1987), pp. 730-757. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28198711%29130%3A5%3C730%3ACRVIDC%3E2.0.CO%3B2-S

Exploitation Ecosystems in Gradients of Primary Productivity Lauri Oksanen; Stephen D. Fretwell; Joseph Arruda; Pekka Niemela The American Naturalist, Vol. 118, No. 2. (Aug., 1981), pp. 240-261. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28198108%29118%3A2%3C240%3AEEIGOP%3E2.0.CO%3B2-2

Food Web Complexity and Species Diversity Robert T. Paine The American Naturalist, Vol. 100, No. 910. (Jan. - Feb., 1966), pp. 65-75. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28196601%2F02%29100%3A910%3C65%3AFWCASD%3E2.0.CO%3B2-D

The Effects of Flow on Polyp-Level Prey Capture in an Octocoral, Alcyonium siderium Mark R. Patterson Biological Bulletin, Vol. 180, No. 1. (Feb., 1991), pp. 93-102. Stable URL: http://links.jstor.org/sici?sici=0006-3185%28199102%29180%3A1%3C93%3ATEOFOP%3E2.0.CO%3B2-9

A Mass Transfer Explanation of Metabolic Scaling Relations in Some Aquatic Invertebrates and Algae Mark R. Patterson Science, New Series, Vol. 255, No. 5050. (Mar. 13, 1992), pp. 1421-1423. Stable URL: http://links.jstor.org/sici?sici=0036-8075%2819920313%293%3A255%3A5050%3C1421%3AAMTEOM%3E2.0.CO%3B2-L

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In Situ Measurements of Flow Effects on Primary Production and Dark Respiration in Reef Corals Mark R. Patterson; Kenneth P. Sebens; R. Randolph Olson Limnology and Oceanography, Vol. 36, No. 5. (Jul., 1991), pp. 936-948. Stable URL: http://links.jstor.org/sici?sici=0024-3590%28199107%2936%3A5%3C936%3AISMOFE%3E2.0.CO%3B2-N

Hydrodynamic Facilitation of Gregarious Settlement of a Reef-Building Tube Worm Joseph R. Pawlik; Cheryl Ann Butman; Victoria R. Starczak Science, New Series, Vol. 251, No. 4992. (Jan. 25, 1991), pp. 421-424. Stable URL: http://links.jstor.org/sici?sici=0036-8075%2819910125%293%3A251%3A4992%3C421%3AHFOGSO%3E2.0.CO%3B2-N

Trophic Interactions in Temperate Lake Ecosystems: A Test of Food Chain Theory Lennart Persson; Sebastian Diehl; Lars Johansson; Gunnar Andersson; Stellan F. Hamrin The American Naturalist, Vol. 140, No. 1. (Jul., 1992), pp. 59-84. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28199207%29140%3A1%3C59%3ATIITLE%3E2.0.CO%3B2-H

Effects of Fish in River Food Webs Mary E. Power Science, New Series, Vol. 250, No. 4982. (Nov. 9, 1990), pp. 811-814. Stable URL: http://links.jstor.org/sici?sici=0036-8075%2819901109%293%3A250%3A4982%3C811%3AEOFIRF%3E2.0.CO%3B2-R

Depth Distributions of Armored Catfish: Predator-Induced Resource Avoidance? Mary E. Power Ecology, Vol. 65, No. 2. (Apr., 1984), pp. 523-528. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28198404%2965%3A2%3C523%3ADDOACP%3E2.0.CO%3B2-7

Hydraulic Food-Chain Models Mary E. Power; Adrian Sun; Gary Parker; William E. Dietrich; J. Timothy Wootton BioScience, Vol. 45, No. 3, Ecology of Large Rivers. (Mar., 1995), pp. 159-167. Stable URL: http://links.jstor.org/sici?sici=0006-3568%28199503%2945%3A3%3C159%3AHFM%3E2.0.CO%3B2-P

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Modes of Feeding in Aggregations of Barnacles and the Shape of Aggregations Julie Pullen; Michael LaBarbera Biological Bulletin, Vol. 181, No. 3. (Dec., 1991), pp. 442-452. Stable URL: http://links.jstor.org/sici?sici=0006-3185%28199112%29181%3A3%3C442%3AMOFIAO%3E2.0.CO%3B2-T

Recruitment Dynamics in Complex Life Cycles Jonathan Roughgarden; Steven Gaines; Hugh Possingham Science, New Series, Vol. 241, No. 4872. (Sep. 16, 1988), pp. 1460-1466. Stable URL: http://links.jstor.org/sici?sici=0036-8075%2819880916%293%3A241%3A4872%3C1460%3ARDICLC%3E2.0.CO%3B2-Y

Factors Regulating Phytoplankton Production and Standing Crop in the World's Freshwaters D. W. Schindler Limnology and Oceanography, Vol. 23, No. 3. (May, 1978), pp. 478-486. Stable URL: http://links.jstor.org/sici?sici=0024-3590%28197805%2923%3A3%3C478%3AFRPPAS%3E2.0.CO%3B2-9

Ecological Energy Relationships at the Population Level L. B. Slobodkin The American Naturalist, Vol. 94, No. 876. (May - Jun., 1960), pp. 213-236. Stable URL: http://links.jstor.org/sici?sici=0003-0147%28196005%2F06%2994%3A876%3C213%3AEERATP%3E2.0.CO%3B2-R

Plaster Standards to Measure Water Motion T. Lewis Thompson; Edward P. Glenn Limnology and Oceanography, Vol. 39, No. 7. (Nov., 1994), pp. 1768-1779. Stable URL: http://links.jstor.org/sici?sici=0024-3590%28199411%2939%3A7%3C1768%3APSTMWM%3E2.0.CO%3B2-Z

Trophic Cascades and Phytoplankton Community Structure Michael J. Vanni; David L. Findlay Ecology, Vol. 71, No. 3. (Jun., 1990), pp. 921-937. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199006%2971%3A3%3C921%3ATCAPCS%3E2.0.CO%3B2-V

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Nutrient Recycling and Herbivory as Mechanisms in the "Top-Down" Effect of Fish on Algae in Lakes Michael J. Vanni; Craig D. Layne Ecology, Vol. 78, No. 1. (Jan., 1997), pp. 21-40. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199701%2978%3A1%3C21%3ANRAHAM%3E2.0.CO%3B2-7

Life and Death in Moving Fluids: Hydrodynamic Effects on Chemosensory-Mediated Predation Marc J. Weissburg; Richard K. Zimmer-Faust Ecology, Vol. 74, No. 5. (Jul., 1993), pp. 1428-1443. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199307%2974%3A5%3C1428%3ALADIMF%3E2.0.CO%3B2-A

Effects of Disturbance on River Food Webs J. Timothy Wootton; Michael S. Parker; Mary E. Power Science, New Series, Vol. 273, No. 5281. (Sep. 13, 1996), pp. 1558-1561. Stable URL: http://links.jstor.org/sici?sici=0036-8075%2819960913%293%3A273%3A5281%3C1558%3AEODORF%3E2.0.CO%3B2-9