Algal Zonation in the New England Rocky Intertidal Community: An Experimental Analysis

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<ul><li><p>Algal Zonation in the New England Rocky Intertidal Community: An Experimental AnalysisAuthor(s): Jane LubchencoSource: Ecology, Vol. 61, No. 2 (Apr., 1980), pp. 333-344Published by: Ecological Society of AmericaStable URL: http://www.jstor.org/stable/1935192 .Accessed: 25/06/2014 06:39</p><p>Your use of the JSTOR archive indicates your acceptance of the Terms &amp; Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp</p><p> .JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact support@jstor.org.</p><p> .</p><p>Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology.</p><p>http://www.jstor.org </p><p>This content downloaded from 37.191.214.157 on Wed, 25 Jun 2014 06:39:36 AMAll use subject to JSTOR Terms and Conditions</p><p>http://www.jstor.org/action/showPublisher?publisherCode=esahttp://www.jstor.org/stable/1935192?origin=JSTOR-pdfhttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsp</p></li><li><p>Ecology, 61(2), 1980, pp. 333-344 ? 1980 by the Ecological Society of America </p><p>ALGAL ZONATION IN THE NEW ENGLAND ROCKY INTERTIDAL COMMUNITY: AN </p><p>EXPERIMENTAL ANALYSIS' </p><p>JANE LUBCHENCO2 Zoology Department, Oregon State University, Corvallis, Oregon 97331 USA </p><p>Abstract. Zonation patterns of plants, including marine algae, have commonly been attributed solely and directly to physical factors. Experimental investigations of the factors affecting zonation of macroscopic, benthic algae in the New England rocky intertidal region demonstrate that biological factors set the lower limits of these plants. The mid zone at all but very exposed sites is usually dominated by brown fucoid algae. These plants are virtually absent in the low zone, which is domi- nated by the red alga Chondrus crispus (Irish moss). Total removal of Chondrus (including the prostrate holdfast) results in establishment of Fucus vesiculosus or F. distichus ssp. edendatus in the low zone. Fucus grows faster in the low than in the mid zone, appears healthy, and reproduces. Thus competition from Chondrus sets the lower limit of Fucus, not changes in light intensity or immersion time, per se, as previously assumed. If herbivores (primarily the periwinkle snail Littorina littorea) are absent where Chondrus is removed, Fucus can settle very densely and occupy 100% of the space. If herbivores are present, Fucus colonizes, but less abundantly. Thus competition is the primary determinant of the zonation pattern (affecting presence or absence) and herbivory is of secondary importance (affecting abundance). </p><p>Other experimental evidence suggests that the upper limit of Chondrus is determined by desic- cation. The lower limit of Chondrus has not been investigated except where a sharp lower limit exists at the low intertidal-shallow subtidal interface. Experiments demonstrate that this is due to the grazing by sea urchins (Strongylocentrotus droebachiensis) where they are locally abundant. Normally, Chon- drus extends well into the subtidal region. </p><p>These results parallel experimental studies of animal zonation in rocky intertidal regions in which biotic factors also set lower bathymetric limits. It is suggested that biogeographic ranges of some species may be similarly affected by biotic factors. </p><p>Key words: algae; Chondrus; community structure; competition; Fucus; herbivory; New En- gland; niche width; zonation. </p><p>INTRODUCTION </p><p>Zonation of plants and animals along environmental gradients is a universal phenomenon. Particularly striking are zonation patterns along rocky intertidal shores. Until the last two decades, studies of rocky intertidal plant and animal zonation were largely de- scriptive (e.g., Stephenson and Stephenson 1972). The correlations of the zones with changes in emergence time, light quality or intensity were assumed to be causal (Gail 1918, Coleman 1933, Hewatt 1937, Doty 1946, Southward 1958, Lewis 1964, Newell 1970; but see Underwood 1978). Beginning with Connell's (1961a, b) studies of factors affecting zonation of bar- nacles, experimental investigation of rocky intertidal animals has demonstrated unequivocally that lower limits of these organisms are usually determined by biological interactions, either predation or competition (Connell 1972). Despite the parallels between sessile animals and plants, and suggestions by numerous workers that competition and herbivory probably af- fect lower limits of algae (Chapman 1957, Southward 1958, Kitching and Ebling 1961, Lewis 1964, Jones and Kain 1967, Vadas 1968, Connell 1972, Chapman 1973, </p><p>1 Manuscript received 4 April 1979; accepted 10 June 1979. 2 J have published previously as Jane Lubchenco Menge. </p><p>1974, Hruby 1976, and Schonbeck and Norton 1978), zonation of marine plants is still often assumed to be solely a function of physical factors: light intensity, light quality, emergence time, submergence time, etc. </p><p>Studies of factors affecting zonation of algae may be interesting for a number of reasons. First, there are strong parallels between studies of bathymetric zo- nation in marine plants and studies of altitudinal, soil moisture, and soil type zonation in terrestrial vegeta- tion. Both have remained largely descriptive (with a few notable exceptions, e.g., Vadas 1968, Dayton 1975, Schonbeck and Norton 1978, and Hruby 1976 for marine algae and Sharitz and McCormick 1973 for terrestrial plants). The majority of terrestrial zonation investigations assume that some abiotic factor that changes along the gradient causes the species replacement patterns because species are specialized to distinct parts of the gradient. Studies are then initiated to isolate the factor or group of fac- tors that best correlates with the vegetation changes. For example, the striking zonation of tree species along an altitudinal gradient in the Green Mountains of Vermont is suggested to be caused by a climatic discontinuity resulting in a sudden change in the num- ber of frost-free days and the amount of moisture and hoar frost present (Siccama 1974). This conclusion is </p><p>This content downloaded from 37.191.214.157 on Wed, 25 Jun 2014 06:39:36 AMAll use subject to JSTOR Terms and Conditions</p><p>http://www.jstor.org/page/info/about/policies/terms.jsp</p></li><li><p>334 JANE LUBCHENCO Ecology, Vol. 61, No. 2 </p><p>remarkably similar to the suggestion that zonation of marine algae is caused by discontinuities in submer- gence times, or so-called "critical tide factors" due to the pecularities of tidal cycles (Coleman 1933, Doty 1946). Given the similarities between these marine and terrestrial patterns perhaps experimental investigation of the processes causing the patterns in one system (the rocky intertidal community) may shed light on the potential mechanisms involved in the other. </p><p>Secondly, experimental studies of zonation patterns may contribute to our understanding of niche widths. Some algae have broad vertical distributions, while others occur only in a narrow band. Factors affecting the widths of the zones are amenable to experimental investigation. </p><p>Thirdly, zonation occurs on a biogeographic scale as well as on a local one. A better understanding of the mechanisms affecting local patterns of distribution and abundance may suggest what factors might cause biogeographic species replacement patterns. </p><p>This paper reports an experimental investigation of factors affecting zonation of the dominant macro- scopic algae in the New England rocky intertidal com- munity. Experiments were designed to evaluate the importance of competition, herbivory, and physical and chemical factors in determining the upper and lower limits of the plants. In particular, the lower limit of the mid zone brown algae Fucus vesiculosus L. and F. distichus L. ssp. edentatus (De la Pyl.) Powell (rockweeds) and the upper and lower limits of the low zone red alga Chondrus crispus Stackhouse (Irish moss) were examined. Experiments were designed based on the wealth of the information available on the biology and ecology of Chondrus (e.g., reviews in Harvey and McLachlan 1973, Prince and Kingsbury 1973a, b, Mathieson and Burns 1975). </p><p>STUDY SITES AND METHODS </p><p>Field observations and experiments were carried out at four main sites along the New England shores. Areas inaccessible to most human disturbance were chosen which represented the range of physical and biological conditions existing along these rocky shores. Detailed descriptions of the sites can be found in J. L. Menge (1975) and B. Menge (1976). The four sites (all in USA) fall along a wave exposure gradient. Ranging from least to most exposed, they are Canoe Beach Cove, Nahant, Massachusetts (42025'N, 70055'W), Grindstone Neck, Maine (44010'N, 6801'W), Chamberlain, Maine (43056'N, 69054'W) and Pema- quid Point (same coordinates as Chamberlain). These areas are stretches of continuous rock (usually granite or basalt) with a minimum of cobbles. </p><p>Basic information on the distribution and abundance of macroscopic plants and animals was obtained every 2-3 mo at each study area using horizontal transects (described more fully in J. L. Menge 1975 and B. </p><p>Menge 1976). Estimates of percent cover of organisms were obtained with a ?/4-M2 plexiglass quadrat which had 100 dots plotted from randomly generated coor- dinates. Estimates of densities of animals were ob- tained by enumerating organisms present within a 1/4- m2 quadrat. </p><p>Space occupiers (plants and sessile animals) exist in three tiers: primary (attached to the substratum), sec- ondary (understory), and tertiary (canopy). The most abundant canopy and understory organisms are three species of fucoid algae and Chondrus, respectively. The discoid holdfasts of the fucoids occupy a very small amount of primary space. Their thalli or upright portions pass through understory space and branch out to occupy extensive amounts of canopy space. The holdfast of Chondrus forms an often extensive, pros- trate red crust on primary space, while the thallus oc- cupies understory space as a shrubby mat. In this pa- per, "upright Chondrus" or just "Chondrus" refers only to the erect thallus, while "Chondrus crust" re- fers only to the prostrate holdfast. </p><p>The effect of Chondrus crispus on the lower limit of Fucus vesiculosus or F. distichus ssp. edentatus was investigated by removing Chondrus from flat sub- strata 1.5-3 m2 in area in the low zone and monitoring subsequent recolonization patterns. In experiments designated "removal of thallus only," the upright or erect portion of Chondrus was scraped off the rocks using putty knives and paint scrapers. This technique leaves the prostrate, encrusting holdfast of Chondrus attached to the rock and is designed to mimic natural removals of Chondrus thallus that occur during winter storms (Lubchenco and Menge 1978). In the experi- ments designated "removal of thallus and crust," the thallus was removed by scraping (as above) and then the crust was removed by burning repeatedly with a propane torch and scraping and brushing vigorously. One exception was an experiment at Chamberlain in which a "removal of thallus only" experiment func- tionally became a "removal of thallus and crust" when the barnacle Balanus balanoides settled very abun- dantly on and completely covered the Chondrus crust. Chondrus' holdfast is exceedingly tenacious and can be removed only with great difficulty. In replicate ex- periments done later, sandblasting was used to remove the crust and proved to be a superior technique to burning. Total thallus and crust removals rarely occur in New England, but may occasionally be caused by limpet grazing and scouring by boulders, cobbles, and ice. </p><p>The angular transformation has been applied to per- cent cover data for statistical analyses (Sokal and Rohlf 1969). In no case did transformations alter the outcome of a statistical test. On figures with mean +95% confidence intervals, percent cover data have been plotted in degrees, then the ordinate labels trans- formed back into percent cover for easier interpreta- </p><p>This content downloaded from 37.191.214.157 on Wed, 25 Jun 2014 06:39:36 AMAll use subject to JSTOR Terms and Conditions</p><p>http://www.jstor.org/page/info/about/policies/terms.jsp</p></li><li><p>April 1980 ALGAL ZONATION 335 </p><p>TABLE 1. Percent cover of the most abundant space occupiers in New England rocky intertidal zones, summer 1974, at four areas along a wave exposure gradient.*t </p><p>Sites arranged from least to most exposed to wave action (left to right) </p><p>Canoe Beach Grindstone Pemaquid Cove Neck Chamberlain Point </p><p>High Zone </p><p>Balanus balanoides 23 ? 8 63 ? 17 93 ? 2 85 ? 7 </p><p>10 Mytilus edulis 13 8 9 12 4 3 5 3 Crustst 43 ? 30 0 0 0 Bare? 17 11 28 + 15 3 ?+3 10 ?+7 </p><p>20 Fucus spiralis 5 ? 7 0 0 0 Ascophyllum nodosum 66 ? 23 0 0 0 </p><p>Mid Zone </p><p>B. balanoides 6 4 1 2 7 2 8 4 </p><p>10 M. edulis 8 7 26 19 66 9 64 13 Crusts 37 29 38 26 0 18 22 Bare 33 25 15 16 3 3 2 3 </p><p>Chondrus crispus 1 ? 1 0 3 ? 2 0 20 Ephemeral algae 0 0 12 ? 8 8 + 9 </p><p>Gigartina stellate 0 0 3 ? 3 0 </p><p>A. nodosum 88 6 0 0 0 30 F. vesiculosus 5 ? 6 78 + 21 0 0 </p><p>F. distichus 0 0 67 + 12 3 3 </p><p>Low Zone </p><p>B. balanoides 0 0 0 37 ? 23 M. edulis 1 2 40 26 52 34 58 21 </p><p>10 Crusts 40 32 11 13 2 2 0 Bare 3 4 8 12 3 3 0 C. c. holdfasts 33 ? 13 24 ? 24 43 ? 34 0 </p><p>20 C. crispus 87 5 68 20 61 28 0 Ephemeral algae 0 0 0 14 ? 14 </p><p>30 F. distichus 0 0 1 ?+3 14 14 Alaria esculenta 0 0 0 44 ? 17 </p><p>* Mean and 95% confidence interval of 10 quadrats in each zone. t Any species occupying 10% of the space is included. 10, 20, and 30 indicate whether the species occupies primary, </p><p>secondary (understory), or tertiary (canopy) space. t Crusts include Verrucaria ericksenii, V. mucosa, Ralfsia verrucosa, R. clavata, Hildenbrandia prototypus, Petrocelis </p><p>middendorfii, Lithothamnium spp., Clathromorphum sp., Phymatolithon sp., and various unidentified red and brown crusts. ? Bare indicates no visible plants or animals present. </p><p>tion. Percent covers of organisms in a single cage and in tables are not transformed. </p><p>COMMUNITY STRUCTURE: ZONATION PATTERNS </p><p>The type and amount of algae present along New England rocky shores vary along two principal en- vironmental gradients, a vertical tidal (depth) and a wave exposure gradient. The striking zonation of plants along the bathymetric gradient has been de- scribed in detail for New England shores (Pearse 1913, Johnson and Skutch 1928, Stephenson and Stephenson 1954, Lamb and Zimmerman 1964, Menge 1976, Lub- chenco and Menge 1978, Mathieson, Hehre and Reyn- olds, in press, and Mathieson, Reynolds and Hehre, in press), as well as for the eastern coasts of the Northern Atlantic where many of the same species occur (references in Chapman 1957, Lewis 1964, Ste- phenson and Stephenson 1972). At areas that range </p><p>from protected to moderately exposed...</p></li></ul>