Peter S. Petraitis and Nicholas Vidargas. 2002. Marine intertidal organisms found in experimental clearings on sheltered shores in the Gulf of Maine, USA. Ecology 87:796.
INTRODUCTION
Different communities often occur in seemingly similar environments, and theorists have explained the existence of these alternative states with plausible mathematical models of species interactions that have several equilibrium points (Lewontin 1969, Noy-Meir 1975, May 1977, Petraitis and Dudgeon 2004a). In these models, the switch between different communities can occur at a critical threshold with a step-like transition in species composition (May 1977, Knowlton 1992). This implies that small environmental perturbations can produce large, discontinuous and irreversible changes in communities. Knowlton (1992) also suggests the critical threshold for a group of species will be correlated with the scale of the perturbation because large events are more likely to cause dramatic shifts in species composition or in interactions among species, which are needed to move the assemblage across the threshold.
It has been suggested that mussel beds and seaweed stands, which are commonly found on sheltered intertidal shores of New England, are alternative community states (Petraitis and Latham 1999). Sheltered shores in the Gulf of Maine are dominated by two distinct assemblages. The first is characterized by the canopy-forming fucoid seaweeds, (Ascophyllum nodosum and Fucus vesiculosus) and the other by beds of barnacles (Semibalanus balanoides) and mussels (Mytilus edulis). Infrequent ice scour events initiate the formation of these two communities in sheltered bays by opening up new clearings, and allowing divergent successional events to pave the way for alternative community states. This is most likely to occur in large clearings where much of a patch may lie beyond the edge effects of the surrounding community, and thus recruits, which arrive in opportunistic fashion, may have sufficient time and space to become established and develop into a divergent assemblage. Once started, a divergent assemblage may be maintained by positive feedbacks and ecosystem engineering within the community (Jones et al. 1994).
Experimental clearings of different sizes were established in A. nodosum stands to test this hypothesis in 1996 at 12 sites on Swan's Island, Maine. The set of clearings at each site included four clearings (1, 2, 4, and 8 m in diameter), and a control plot that was not cleared for a total of 60 plots (Petraitis and Dudgeon 1999). Analysis of data on densities and percent cover of the most common marine organisms from 1996 to 2002 shows that succession depend on clearing size and not on site-specific differences (Petraitis et al. 2003; Petraitis and Dudgeon 2005).
METADATA
CLASS I. DATA SET DESCRIPTORS
A. Data set identity: Abundance and Percent Cover of Marine Organisms on Sheltered Intertidal Shores in the Gulf of Maine
B. Data set identification code: Succession_sampling_96-02_data.txt
C. Data set description
Principal Investigator: Peter S. Petraitis, Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018 USA
Abstract: Experimental clearings in macroalgal (Ascophyllum nodosum) stands were established in 1996 to determine if mussel beds and macroalgal stands on protected intertidal shores of New England represent alternative community states. Circular clearings 1, 2, 4, and 8 m in diameter were made in Ascophyllum stands at 12 sites on Swan’s Island, Maine. Each site also had an uncleared control. The sites are nested within four bays. The purpose of this data set is to provide access to data on densities and percent cover in the 60 experimental plots from 1996 to 2002. Data include densities of mussels (Mytilus edulis and Modiolus modiolus), an herbivorous limpet (Tectura testudinalis), herbivorous snails (Littorina littorea, Littorina obtusata), a predatory snail (Nucella lapillus), a barnacle (Semibalanus balanoides), and fucoid algae (Ascophyllum nodosum and Fucus vesiculosus), and percent cover by mussels, barnacles, and fucoids. Published analyses of the data set show that succession depends on clearing size and not on site-specific differences.
D. Key words: alternative stable states; community ecology; Gulf of Maine; LTREB data; marine algae; marine invertebrates; mussels; rocky intertidal shores; succession
CLASS II. RESEARCH ORIGIN DESCRIPTORS
A. Overall project description
Identity: Abundance and Percent Cover of Marine Organisms on Sheltered Intertidal Shores in the Gulf of Maine
Originator: Peter Petraitis, Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018.
Period of Study: 1996 – 2002 (ongoing)
Objectives: To use successional changes in experimental clearings to test hypothesis concerning alternative community states.
Abstract: Same as above.
Sources of funding: All data collection has been supported by NSF (OCE 95-29564 and DEB LTREB 03-14980).
B. Specific subproject description
Site description: Experimental clearings were established at 12 replicate sites in four bays on Swan’s Island, Maine, USA with three sites nested in each bay. The bays are Mackerel Cove and Seal Cove, which are on the north side of the island, and Burnt Cove Harbor and Toothacker Cove, which are on the south side of the island. See Dudgeon and Petraitis (2001, Fig. 2) for map with locations of sites and bays.
Site type: The sites are in the mid intertidal zones of sheltered bays, and experimental clearings are between 0.3 and 1.0 m above Mean Low Water.
Geography: Swans Island is located at 44°10’ N, 68°25’ W in the Gulf of Maine, USA. The island is approximately 36 km2 in area with a highly irregular coastline.
Habitat: The mid intertidal shores are protected from wave surge and dominated by the rockweed A. nodosum.
Geology: The shoreline surfaces at the different sites are a mixture of granite and basalt outcrops, boulder fields, and some muddy patches in the most protected sites.
Watersheds/hydrology: The exposure to waves at these sites ranges from extremely protected to moderately protected.
Site history: N/A
Climate: Coastal northeastern USA
Experimental design: At each site, four circular experimental plots, which were 1, 2, 4, and 8 m in diameter clearings, and a control plot, which was not cleared, were established. It was not possible to make the clearings during the winter because of limited access and dangerous conditions due to snow and ice. Thus plots were originally scraped between 22 June and 17 August 1996 and then re-scraped between 6 and 10 February 1997 to mimic winter ice events. Scraping was done using paint scrapers and removed all A. nodosum. Center of plots were marked with two stainless steel reference bolts which were anchored in drilled holes. Plots were left unmanipulated thereafter. Plots within a site are separated by at least 10 m, and sites within bays by at least 750 m. The size range of the experimental clearings is within the normal range of major ice scour events, which occur infrequently on Swan’s Island. For example, ice removed an average of 13% of the A. nodosum cover from the 12 experimental sites during the winter of 2002–2003 (range: 0–39%, P. S. Petraitis, unpublished data). At the three sites with the greatest damage, the average area of naturally created clearings was 25.4 m2 (range: 2.4–78.5 m2, n = 5 clearings per site). The experimental clearings are 0.8–50.3 m2 in area.
Design characteristics: The experimental set-up is a partially nested design with clearing size fully crossed with bays and sites, and sites nested within bays. Sites and bays are considered random effects and clearing size is a fixed treatment effect. The design is fully discussed in Petraitis and Dudgeon (2004b).
Sampling methods: Abundance and percent cover of the most common species have been sampled at least once a year since 1996. This data set contains data from summer 1996, spring and summer 1997, spring and summer 1998, summer 1999, spring and summer 2000, and summers of 2001 and 2002. Percent cover and densities were not sampled for all species on all dates. Uncleared control plots were not sampled in 1996. Spring samples are taken between mid-May and mid-June (median date: 15 June), and summer samples are taken between late June and mid August (median date: 7 July).
Mussels (Mytilus edulis and Modiolus modiolus) and gastropods (Tectura testudinalis, Littorina littorea, Littorina obtusata and Nucella lapillus) were counted using five 50 × 50 cm quadrats per plot. Fucoid algae (Ascophyllum nodosum and Fucus vesiculosus) were counted in the five 10 × 10 cm squares on the diagonal of each 50 × 50 cm quadrat because of the large number of recruits. The barnacle Semibalanus balanoides was counted in two 2 × 2 cm squares per 50 × 50 cm quadrat except in 1998 when one square per quadrat was used. Barnacle counts were divided into the current year’s recruits and older individuals, which could be distinguished by differences in size and color.
Percent cover data were collected for fucoids, barnacles and mussels. Cover was estimated by counting the most common space occupier in the twenty-five 10 × 10 cm squares per 50 × 50 cm quadrat. Percent cover was thus calculated on the basis of 125 counts of presence/absence (25 squares per quadrat × 5 quadrats per plot). Ascophyllum nodosum was considered the normal canopy species and was counted first (range 0–100%) and then was pulled back to reveal the common understory species (Fucus vesiculosus, Semibalanus balanoides and Mytilus edulis), which were then counted. The category of “Bare” includes all other species, which were rare, and bare space. Understory species and bare space sums to 100%. Quadrats were grouped in the center of each clearing, and the sampling in the uncleared controls was done within 2 m of the reference bolts.
Taxonomy and systematics: Names for algae follow usage in Taylor (1957) and for invertebrates follow usage in Gosner (1971) except for the limpet, which has been revised from Acmaea testudinalis to Tectura testudinalis (current name retrieved 30 November 2005 from the Integrated Taxonomic Information System; www.itis.usda.gov; taxonomic serial number: 204945).
Permit history: Clearings were made under permit from the State of Maine’s Department of Marine Resources.
Legal/organizational requirements: None.
Project personnel: Peter Petraitis, Steve Dudgeon, Erika Carlson Rhile, Nick Vidargas
CLASS III. DATA SET STATUS AND ACCESSIBILITY
A. Status
Latest update: The data set spans the period of 1996–2002. Data collection is ongoing through the present and will be added as collected and verified.
Latest Archive date: 4 June 2005
Metadata status: The metadata are complete and up to date.
Data verification: Original data entries in Excel files are checked prior to calculation of averages. Averages are screened for outliers, which are then re-checked by comparing hand-written entries on original data books against entries in Excel files. Averages for percent cover are also checked to see if sum of understory species (barnacles, mussels, Fucus, and bare surface) sums to 100%.
B. Accessibility
Storage location and medium: (Ecological Society of America data archives [Ecological Archives link], URL published in each issue of its journals). Original data file exists on author’s personal computer and on CD in MSExcel format.
Contact person: Peter Petraitis, email: ppetrait@sas.upenn.edu, Tel. 215.898.4207, Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018, USA
Copyright restrictions: None.
Proprietary restrictions: None.
Costs: None.
CLASS IV. DATA STRUCTURAL DESCRIPTORS
A. Data Set File
Identity: Succession_sampling_96-02_data.txt
Size: 588 records, not including header row.
Format and storage mode: ASCII text, tab delimited. No compression scheme was used.
Header information: See variable names in Section B.
Alphanumeric attributes: Mixed.
Special characters/fields: Missing data denoted as -999.9
Authentication procedures: The sums of the numeric columns are:
LL = 24767.8, LO = 2072.4, NL = 899.1, TT = 419.1, FV = -94154.4, AN =-115350.4, ME = 5681.0, MM = 134.9, OSB = 465.9,
NSB = -45456.8, PSB = -283024.6, PFV = -162003.2, PAN = -283568.4, PME = -291671.8, PBR = -278740.3
B. Variable information
Latitude and Longitude: Latitude and longitude of reference bolts in experimental plots in degrees and decimal minutes. Positions determined in summer 2005 using a handheld GPS-WAAS receiver (Lowrance iFinder) with accuracy within 2 m.
Date: Date on which plots were sampled.
Yr_code: Yr_codes from 1997 to 2002 were used as “Date” variable in univariate analyses in Petraitis and Dudgeon (2005). Yr_code gives year of sampling followed by “a” for spring samples and “b” for summer samples.
Bay: Names of bays are consistent with names on U.S. Geological Survey 7.5 minute topographic quadrangle for Swans Island. Codes for bay names are: BC = Burnt Coat Harbor, MC = Mackerel Cove, SC = Seal Cove, TC = Toothacker Cove.
Site: Site names were assigned based on either names that appear on USGS maps (Long Cove and Hen for Hen Island), or individuals who provided access to the shore or live near a site (McD, Bob, Dick, Basil, and Grace), or local names that are not on maps (Ledges, Mill Pond, and Little House Cove), or were arbitrary choices (July 4th for the date that the clearings were made at the site, and Pole 24 for location of a telephone line pole). Codes for site names are the same as shown on the map in Dudgeon and Petraitis (2001, Fig. 2) and are: BO = Bob, BS = Basil, DK = Dick, GR = Grace, HN = Hen, JL = July 4th, LC = Long Cove, LD = Ledges, LH = Little House Cove, MD = McD, MP = Mill Pond, and PL = Pole 24.
Size: Diameter of experimental clearings. Uncleared control plots are given as 0.
Numeric variables: Variables of interest are densities and percent cover per plot, and so the data set contains the average densities and average percent cover per plot based on the quadrats per plot. Note that counts per quadrat may not be independent because randomly placed quadrats overlapped in some plots; however the average of a plot, as long as the observations per plot were taken at random, provides an unbiased estimate of the parametric mean of the plot (Mood and Graybill, 1963:142–153). There is the potential for problems with precision of the estimates because differences in the spatial spread and overlap of quadrats could have caused heterogeneity of variances, but we found no evidence for this.
Variable name |
Variable definition |
Units |
Storage type |
Range numeric values |
Missing value codes |
Latitude |
Latitude |
Degrees decimal minutes |
Character |
N/A |
N/A |
Longitude |
Longitude |
Degrees decimal minutes |
Character |
N/A |
N/A |
Date |
Sampling date |
N/A |
Character |
N/A |
N/A |
Yr_code |
Year |
N/A |
Character |
N/A |
N/A |
Bay |
Bay |
N/A |
Character |
N/A |
N/A |
Site |
Site |
N/A |
Character |
N/A |
N/A |
Size |
Diameter of clearing |
N/A |
Character |
N/A |
N/A |
LL |
Average abundance Littorina littorea |
No./0.25 m2 |
Floating Point |
0.6 – 356.0 |
-999.9 |
LO |
Average abundance Littorina obtusata |
No./0.25 m2 |
Floating Point |
0 – 118.6 |
-999.9 |
NL |
Average abundance Nucella lapillus |
No./0.25 m2 |
Floating Point |
0 – 28.0 |
-999.9 |
TT |
Average abundance Tectura testudinalis |
No./0.25 m2 |
Floating Point |
0 – 15.6 |
-999.9 |
FV |
Average abundance Fucus vesiculosus |
No./0.25 m2 |
Floating Point |
0 – 779.0 |
-999.9 |
AN |
Average abundance Ascophyllum nodosum |
No./0.25 m2 |
Floating Point |
0 – 730.0 |
-999.9 |
ME |
Average abundance Mytilus edulis |
No./0.25 m2 |
Floating Point |
0 – 1393.6 |
-999.9 |
MM |
Average abundance Mytilus modiolus |
No./0.25 m2 |
Floating Point |
0 – 17.0 |
-999.9 |
OSB |
Average abundance old Semibalanus balanoides |
No./4 cm2 |
Floating Point |
0 – 8.6 |
-999.9 |
NSB |
Average abundance new S. balanoides |
No./4 cm2 |
Floating Point |
0 – 32.6 |
-999.9 |
PSB |
Average cover S. balanoides |
Percent |
Floating Point |
0 – 100.0 |
-999.9 |
PFV |
Average cover F. vesiculosus |
Percent |
Floating Point |
0 – 100.0 |
-999.9 |
PAN |
Average cover A. nodosum |
Percent |
Floating Point |
0 – 100.0 |
-999.9 |
PME |
Average cover M. edulis |
Percent |
Floating Point |
0 – 30.4 |
-999.9 |
PBR |
Average cover by bare space, other species |
Percent |
Floating Point |
0 – 100.0 |
-999.9 |
CLASS V. SUPPLEMENTAL DESCRIPTORS
A. Data acquisition
Data forms: “Rite in the Rain” field log books.
Location of completed data forms: 325 Leidy Labs, Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018, USA
Data entry/verification procedures: Field crews were supplied with log books to record all data in the field. Data were then entered into a computer in the lab and double-checked. Files are stored on author’s personal computer and on CD as MSExcel files. Log books are held at author’s address.
B. Quality assurance/quality control procedures: See comments in comments on data verification (Class III, Section A), data entry/vertification procedures (Class V, Section A), and computer programs and data processing algorithms (Class V, Section D).
C. Related material: Data on adult mussel mortality (1996, 1999, 2000, 2003, and 2004) and recruitment of barnacles, mussels, and fucoids (1997–2005) in the experimental clearings has been collected but only results prior to 1997 have been published (Petraitis and Dudgeon 1999, Dudgeon and Petraitis 2001).
D. Computer programs and data processing algorithms: Averages were calculated using the “average” function in MSExcel. Checks for outliers in numeric variables used sort function in MSExcel to examine extreme values and used “if” statements to check summations of percent cover.
E. Archiving: N/A
F. Publications using the data set:
Petraitis, P.S., E.C. Rhile, E.C., and S.R. Dudgeon. 2003. Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities. Journal of Experimental Marine Biology and Ecology 293:217–236.
Petraitis, P.S., and S.R. Dudgeon. 2005. Divergent succession and implications for alternative states on rocky intertidal shores. Journal of Experimental Marine Biology and Ecology 326:14–26.
G. Publications using the same sites:
Dudgeon, S.R., J.E. Kübler, W.A. Wright, R.L. Vadas, Sr., and P.S. Petraitis. 2001. Natural variability in zygote dispersal of Ascophyllum nodosum at small spatial scales. Functional Ecology 15:595–604.
Dudgeon, S.R., and P.S. Petraitis. 2001. Scale-dependent recruitment and divergence of intertidal communities. Ecology 82:991–1006
Dudgeon S. R., and P.S. Petraitis. 2005. First year demography of a foundation species, Ascophyllum nodosum, and its community implications. Oikos 109:405–415.
Petraitis, P.S., and S.R. Dudgeon. 1999. Experimental evidence for the origin of alternative communities on rocky intertidal shores. Oikos 84:239–245.
H. History of data set usage
Data request history: N/A
Data set update history: Petraitis et al. (2003) and Petraitis and Dudgeon (2005) used earlier versions of this data set. In particular, several minor errors in percent cover data used in Petraitis and Dudgeon (2005) were found and corrected is this updated version.
Review history: N/A
Questions and comments from secondary users: N/A
ACKNOWLEDGMENTS
We thank the residents of Swans Island who supported this research by provided access to the shore across their properties. Research was supported by National Science Foundation grants (OCE 95-29564 and DEB LTREB 03-14980) to P. S. Petraitis.
LITERATURE CITED
Dudgeon, S., and P. S. Petraitis. 2001. Scale-dependent recruitment and divergence of intertidal communities. Ecology 82:991–1006.
Gosner, K. L. 1971. Guide to identification of marine and estuarine invertebrates. John Wiley, New York, New York, USA.
Jones, C. G., J. H. Lawton, and M. Shachak. 1994. Organisms as ecosystem engineers. Oikos 69:373–386.
Knowlton, N. 1992. Thresholds and multiple stable states in coral reef community dynamics. American Zoologist 32:674–682.
Lewontin, R. C. 1969. The meaning of stability, Diversity and stability in ecological systems. Brookhaven National Laboratory, Upton, New York.
May, R. M. 1977. Thresholds and breakpoints in ecosystems with a multiplicity of stable states. Nature 269:471–477.
Mood, A. M., and F. A. Graybill. 1963. Introduction to the theory of statistics. McGraw-Hill, New York, New York, USA.
Noy-Meir, I. 1975. Stability of grazing systems: an application of predator-prey graphs. Journal of Ecology 65:459–481.
Petraitis, P. S., and S. R. Dudgeon. 1999. Experimental evidence for the origin of alternative communities on rocky intertidal shores. Oikos 84:239–245.
Petraitis, P. S., and S. R. Dudgeon. 2004a. Detection of alternative stable states in marine communities. Journal of Experimental Marine Biology and Ecology 300:343–371.
Petraitis, P. S., and S. R. Dudgeon. 2004b. Do alternate stable community states exist in the Gulf of Maine rocky intertidal zone? Comment. Ecology 85:1160–1165.
Petraitis, P. S., and S. R. Dudgeon. 2005. Divergent succession and implications for alternative states on rocky intertidal shores. Journal of Experimental Marine Biology and Ecology 326:14–26.
Petraitis, P. S., and R. E. Latham. 1999. The importance of scale in testing the origins of alternative community states. Ecology 80:429–442.
Petraitis, P. S., E. C. Rhile, and S. R. Dudgeon. 2003. Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities. Journal of Experimental Marine Biology and Ecology 293:217–236.
Taylor, W. R. 1957. Marine algae of the northeastern coast of North America. Second edition. The University of Michigan Press, Ann Arbor, Michigan, USA.