Marine Sanctuaries Conservation Series ONMS-10-05

U.S. Department of Commerce 

National Oceanic and Atmospheric Administration 

National Ocean Service 

Office of Ocean and Coastal Resource Management 

Office of National Marine Sanctuaries December 2010

Dynamics of Hard Substratum Communities 

Inside and Outside of a Fisheries Habitat Closed 

Area in Stellwagen Bank National Marine 

Sanctuary (Gulf of Maine, NW Atlantic) 
About the Marine Sanctuaries Conservation Series 

The National Oceanic and Atmospheric Administration’s National Ocean Service (NOS) 

administers the Office of National Marine Sanctuaries (ONMS). Its mission is to identify, 

designate, protect and manage the ecological, recreational, research, educational, 

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programs is fundamental to marine protected area management. The Marine 

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forum for publication and discussion of the complex issues currently facing the sanctuary 

system. Topics of published reports vary substantially and may include descriptions of 

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scientific research and monitoring projects. The series facilitates integration of natural 

sciences, socioeconomic and cultural sciences, education, and policy development to 

accomplish the diverse needs of NOAA’s resource protection mandate. 
Dynamics of Hard Substratum Communities Inside and 

Outside of a Fisheries Habitat Closed Area in Stellwagen Bank 

National Marine Sanctuary (Gulf of Maine, NW Atlantic) 

Alison Tamsett, Kari B. Heinonen, Peter J. Auster 

Department of Marine Sciences and 

Northeast Undersea Research Technology & Education Center 

University of Connecticut 

Groton, CT 06340 

 

and 

James Lindholm 

California State University Monterey Bay 

Division of Science and Environmental Policy and 

Institute for Applied Marine Ecology 

Seaside, CA 93955 

 Silver Spring, Maryland 

 December 2010 

 U.S. Department of Commerce 

Gary Locke, Secretary 

National Ocean and Atmospheric Administration 

Jane Lubchenco, Ph.D. 

Under Secretary of Commerce for Oceans and Atmosphere 

National Ocean Service 

David Kennedy, Acting Assistant Administrator 

Office of National Marine Sanctuaries 

Daniel J. Basta, Director 
Disclaimer 

Report content does not necessarily reflect the views and policies of the Office of 

National Marine Sanctuaries or the National Oceanic and Atmospheric Administration, 

nor does the mention of trade names or commercial products constitute endorsement or 

recommendation for use. 

Report Availability 

Electronic copies of this report may be downloaded from the Office of National Marine 

Sanctuaries web site at http://sanctuaries.noaa.gov. Hard copies may be available from 

the following address: 

 National Oceanic and Atmospheric Administration

 Office of National Marine Sanctuaries

 SSMC4, N/ORM62 

 1305 East-West Highway 

 Silver Spring, MD 20910 

Cover

A diversity of organisms live on hard substratum surfaces in the deep waters of 

Stellwagen Bank National Marine Sanctuary. 

Suggested Citation

Tamsett, A., K.B. Heinonen, P.J. Auster and J. Linholm. 2010. Dynamics of hard 

substratum communities inside and outside of a fisheries habitat closed area in 

Stellwagen Bank National Marine Sanctuary (Gulf of Maine, NW Atlantic). Marine 

Sanctuaries Conservation Series ONMS-10-05. U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration, Office of National Marine Sanctuaries, Silver 

Spring, MD. 53 pp. 

Contact 

Peter J. Auster, Department of Marine Sciences and Northeast Underwater Research 

Technology & Education Center, University of Connecticut at Avery Point, 1080 

Shennecossett Rd., Groton, CT 06340 USA. Email: peter.auster@uconn.edu
i 

Abstract 

The dynamics of gravel and boulder reef invertebrate community structure in areas inside 

and outside a closed fisheries habitat that overlaps Stellwagen Bank National Marine 

Sanctuary were analyzed based on time-series photographic transects from 1998-2005. 

The effectiveness of this protected area was evaluated based on three common 

predictions that emerge from previous studies of fishing impacts and the performance of 

temperate marine protected areas. We determined if: (1) gravel and boulder habitats have 

similar community composition, (2) community structure diverged between fished and 

unfished sites attributable to chronic fishing impacts, (3) structure forming invertebrates 

increased in abundance within the protected area, and (4) diversity increased within the 

protected area. Overall our results demonstrate that community structure over the seven 

years since closure in 1998 has been dynamic across both habitat types as well as within 

and outside the Western Gulf of Maine Closure (WGOMC) despite a high degree of 

similarity between paired habitat stations at the time of closure. Comparisons of each 

habitat type inside and outside the closure across years in regards to community structure, 

populations of component taxa, and patterns of diversity all demonstrated a response to 

the closure but not in ways that are normally predicted from previous closed areas 

studies. Despite the presence of hard substratum resources in both boulder and gravel 

habitats, community structure was different between habitat types across all years. 

Community structure changed across time both inside and outside the WGOMC 

suggesting, at least to-date, recovery without resilience. While community composition 

tended to be more similar within each station than between each year, the pattern of 

similarity from 2005 transects suggest a greater degree of difference in composition 

between replicates from inside gravel and boulder stations than those paired stations 

outside. This pattern suggests the dominance of local processes, such as predation and 

competition, may be driving community composition inside the closed areas (i.e., 

contributing to greater variation in the distributions of taxa within stations). This is in 

contrast to larger spatial scale disturbance processes, produced either by natural events or 

by fishing activities that dominate at outside stations. Species populations and 

community structure within the closed area have yet to reach any stable configuration. 

Interestingly, structural guilds and population trajectories of component taxa changed 

over time in unpredictable ways. We predicted that structure forming invertebrates 

would increase in abundance over time within the protected area due to elimination of 

fishing gear disturbance and a recovery of erect and emergent fauna. However, only 

encrusting forms at the boulder stations outside the closed area increased significantly 

from 1998 to 2005. Finally, the expected increase in species diversity at stations within 

the WGOMC was not observed by the end of the study period at either gravel or boulder 

stations. Overall the findings indicate that the WGOMC is having a significant impact on 

invertebrate community structure and that the community inside the closure area on both 

boulder and gravel habitats is recovering from chronic fishing gear impacts. However, 

community structure is dynamic and that “recovery” of the seafloor community does not 

necessarily lead to a climax community. 
ii 

Key Words 

Marine protected area, resilience, recovery, disturbance, benthic, fishing, photography 
iii 

Table of Contents 

Topic Page

Abstract ................................................................................................................................ i

Key Words ......................................................................................................................... iii

Table of Contents ............................................................................................................... iii

List of Figures and Tables.................................................................................................. iv

Introduction ......................................................................................................................... 1

Materials and Methods ........................................................................................................ 3

 Sample Collection .......................................................................................................... 3 

 Analysis.......................................................................................................................... 5 

Results ................................................................................................................................. 6 

Discussion ......................................................................................................................... 33

Acknowledgements ........................................................................................................... 36

Literature Cited ................................................................................................................. 36 

Appendix 1: Sampling and data collection details ........................................................... 41

Appendix 2: Details of pairwise comparisons ................................................................. 44

 
iv 

List of Figures and Tables 

Figure/Table Number and Title Page

Figure 1. Location map of sampling stations ..................................................................... 4

Figure 2. MDS of gravel and boulder habitat community composition ............................ 9

Figure 3. MDS of habitat types over time ........................................................................ 10

Figure 4. K dominance curves for gravel and boulder habitat ......................................... 17 

Figure 5. Mean abundance of selected taxa over time ..................................................... 21

Figure 6. Species diversity indices for gravel habitat ...................................................... 30

Figure 7. Species diversity indices for boulder habitat .................................................... 31 

Table 1. ANOSIM analysis between habitat types, years, and closure status ................. 11

Table 2. SIMPER analysis between habitat types ........................................................... 11

Table 3. SIMPER analysis for gravel habitat by closure status ....................................... 11

Table 4. SIMPER analysis for gravel stations by years ................................................... 12

Table 5. SIMPER analysis for boulder habitat by closure status ..................................... 14

Table 6. SIMPER analysis for boulder stations by years ................................................. 15

Table 7. ANOVA for individual taxa and structural groups ............................................ 18

Table 8. Pairwise comparisons summary matrix for taxonomic categories .................... 25 

Table 9. Pairwise comparisons summary matrix for structural groups ........................... 26

Table 10. ANOVA for species diversity indices ............................................................. 32

 
1

Introduction 

The National Marine Sanctuary Act (NMSA) requires that marine sanctuaries are 

managed to allow multiple uses (e.g., fishing), assuming that those activities are 

compatible with the primary goal, which is resource protection. However, there are no 

clear metrics and reference points to trigger management actions (Samhouri et al. 2010) 

and information to guide decision-making is often fragmented and incomplete. The need 

to balance human uses of the marine environment with the conservation of biological 

diversity requires knowledge of how particular activities affect diverse populations and 

communities within particular habitats (Barr 1995). Further, the reauthorization of the 

Magnuson Fishery Conservation and Management Act in 1996 mandated the 

identification and protection of Essential Fish Habitat (EFH) defined as “those waters and 

substrate necessary to fish for spawning, breeding, feeding, or growth to maturity” 

(Schmitten 1999), but like the NMSA, there are no clear reference points for action. One 

management strategy to conserve EFH and biological diversity overall is the use of yearround closed areas (National Research Council 2002). Such areas have been used off the 

northeast United States, both as ad hoc areas of habitat protection when regions were 

closed to reduce fishing mortality on selected species as well as areas closed specifically 

for habitat management (Link et al. 2005, Murawski et al 2005). Gravel and boulder 

habitats are of particular concern as attributes of habitat complexity have been linked to 

survivorship of species of economic importance (Auster et al. 1996, Auster 1998, 

Lindholm et al. 1999, 2001). 

Bottom contact fishing gears have been found to have significant effects on seafloor 

communities and habitats (e.g., Watling and Norse 1998, Auster and Langton 1999, 

National Research Council 2002, Kaiser et al. 2006). In particular, use of mobile gear 

can reduce habitat complexity by removing and damaging emergent fauna, smoothing 

seafloor features (e.g., small patches of piled boulders and sand waves), and removing 

structure producing taxa (e.g., crabs and fish that produce depressions and burrows; 

Auster 1998, Auster et al. 1996, Collie et al. 1997, Collie et al. 2005,). Long-term studies 

to evaluate recovery rates under a variety of conditions, both in terms of gear type and 

community settings, are rare (Auster and Langton 1999, Kaiser et al. 2006). Two metaanalyses of results from available gear impact studies around the globe, across a range of 

gear types and from multiple habitats, suggest recovery times of hard substrate 

communities in temperate ecosystems on the order of 8-10 or more years (Collie et al. 

2000, Kaiser et al. 2006). Empirical studies of gravel habitat communities on the 

Northeast Peak of Georges Bank, subject to strong tidal currents and a well-mixed water 

column, have recovery times in excess of ten years based on time-series monitoring 

(Collie et al. 2005). Such patterns are consistent with our understanding of the life 

histories of many large habitat-forming epifaunal species (e.g., sponges, corals, 

calcareous worms, bryozoa) that tend to be long-lived and ill-adapted to frequent 

disturbance (Auster et al. 1996, Watling and Norse 1998). In this region undisturbed 

deep-water gravel habitats were found to have increased biomass, species richness, and 

species abundance relative to comparable gravel sites in fished areas (Collie et al 2005). 
2

The results of these previous studies indicate that hard-substratum epifaunal communities 

are not likely resilient to disturbances by fishing gear but can recover. The resilience of a 

community can be evaluated by determining if the community: 1) can be disturbed and 

recover to its prior state; 2) is able to resist the disturbance (i.e., remains essentially 

unchanged following a disturbance event); or 3) shifts to another state following a 

disturbance event (Palumbi et al. 2008). Communities that exhibit resilience to 

disturbance will either rapidly return to the pre-disturbed state (recovery) or remain 

fundamentally unchanged following a disturbance (resilient). Communities that are not 

resilient to disturbance will recover to an altered community state (Auster and Langton 

1999). Prior studies indicate that hard-substrate habitats may exhibit some level of 

resilience to disturbance by reaching a state of “recovery” within a particular time period. 

Alternatively, recovery may indicate the community is not resilient to disturbance, and 

instead shifts to an alternative community state. 

 

There are two generally accepted models of community dynamics (see Auster and 

Langton 1999 and references therein). The first is the traditional linear successional 

model where a disturbed community recruits “pioneer” species that alter the local 

environment to a state favorable to recruitment of sequential “intermediate” species, that 

then further modify an environment to be suitable for recruitment and development of a 

sustained “climax” community. The second model is a lottery-based model where shifts 

in the community structure result from competition among species or disturbance events 

that alter the environment to one that favors a different community composition. In such 

a model, the shifts in the community structure are not predictable and do not follow a 

sequential successional pattern. Communities that fit either model may be resilient to 

disturbances. In the successional model, resilient communities that are subjected to 

disturbance may: 1) remain unchanged (i.e., resilient), or 2) revert back to a prior state or 

may “skip” a step in the linear progression to the climax community, but will always 

return to the climax community state over time (i.e., recovery). In the lottery-based 

model, disturbed communities may remain unchanged following disturbance (i.e., 

resilient); or 2) shift to another stable community state (i.e., recovery without resilience). 

To evaluate the model that applies in a community subjected to disturbance a stable state 

must be reached and maintained. 

Stellwagen Bank National Marine Sanctuary (SBNMS) is a multiple use marine protected 

area located in the western Gulf of Maine. The sanctuary contains Stellwagen Bank as 

well as many other high relief topographic features composed of a range of coarse 

substratum (Battista et al. 2006). The eastern side of Stellwagen Bank is isolated from 

sediment sources while sands that drape a gravel pavement on the top of the bank are 

eroding over time. Tidal currents are weak and reach maximum speeds of 20–30 cm s−1

. 

However, the bank lies in the path of strong northeasterly storms and currents generated 

by storm waves in the deep waters of the Gulf of Maine modify the seabed as they pass 

over the bank to depths of 50 – 80 m (Valentine & Schmuck 1995). Internal waves 

impinge on the bank and affect rates of food delivery and invertebrate larvae for 

recruitment at the seafloor (Battista et al. 2006, Haury et al. 1983, Witman et al. 1993). 

 

The Western Gulf of Maine Closure (WGOMC), a fisheries closure implemented in 1998 
3

(Murawski et al. 2005), encompasses approximately 21% of SBNMS and includes gravel 

pavement and boulder reef habitats. Originally the closure prohibited the use of mobile 

fishing gear capable of catching groundfish species such as cod and haddock. In 2004, 

the closure was redesignated as a habitat closure and all mobile, bottom-tending fishing 

gear was prohibited. However, lobster traps, recreational hook-and-line, and mid-water 

trawls are still allowed in the closure area. 

 

Here we report on the dynamics of gravel and boulder reef invertebrate community 

structure inside and outside the WGOMC within SBNMS from 1998 - 2005. Further, we 

evaluate the effectiveness of this marine protected area based on four common issues that 

emerge from previous studies of fishing impacts and the effectiveness of temperate 

marine protected areas (e.g., Auster and Shackell 2000, Auster and Langton 1999, Link et 

al. 2005). In particular, we determine if: (1) gravel and boulder habitats have similar 

community composition, (2) community structure diverges between fished and unfished 

sites attributable to chronic fishing impacts, (3) structure forming invertebrates increase 

in abundance within the protected area, and (4) diversity increases within the protected 

area. Based on the results of these comparisons, we address issues of resilence and their 

fit to models of community dynamics. While time series data will continue to be 

collected at these stations, the results in this report have implications for management of 

biological diversity within SBNMS as well as management of EFH across the Gulf of 

Maine region. 

Materials and Methods 

Sample Collection 

Monitoring of seafloor communities on a near annual basis was performed inside and 

outside the WGOMC in Stellwagen Bank National Marine Sanctuary using underwater 

still and video imaging (Appendix 1, Table 1). Replicate transects at each habitat (i.e., 

gravel and boulder) by fishing treatment (inside and outside the closed area) were 

conducted over a seven year time period from 1998 to 2005. Paired stations for each 

habitat type were chosen at the time of closure (i.e., 1998) based on knowledge of 

seafloor geology from a continuous coverage multibeam sonar map (Valentine et al. 

2003) as well as direct observations made from underwater video of the seafloor . Station 

selection was constrained by minimizing differences in depth and distance to the greatest 

extent possible between stations within each habitat treatment (Figure 1). A minimum of 

three transects composed of twenty photographs each (one photo per minute for the 

duration of a twenty minute transect) were conducted at each station during each year. 

Three different vehicle platforms were used to capture seafloor images over the time 

series (i.e., two different ROVs and a camera sled) but all acquired data in the same 

manner. Photographs were obtained using a down-looking 35mm film camera with an 

electronic flash unit for illumination. Two lasers mounted in parallel (20 cm spacing) 

were used to calibrate images and to maintain camera height at 0.75 m. Area of each 

photograph was nominally 0.39 m2

. The three transect lines were offset by 

approximately 50 m at each station in order to ensure no areas of overlap and 

independence of photographs between transects. Transects were conducted by drift or 
4

active motoring depending on the support vessel. While there was year-to-year variation 

in the exact location of transects, all were conducted within an approximate 0.50 km 

Figure 1. Location of sampling stations in the Stellwagen Bank National Marine Sanctuary. 

Hatched area is Western Gulf of Maine Closure (WGOMC). Boundary in white is Stellwagen Bank 

National Marine Sanctuary. Sampling stations are noted by substrate type. Sand and mud stations 

were evaluated using different methods and will be reported elsewhere. 

radius of the sample station. Invertebrate fauna recorded in the photographs were 

identified to the lowest possible taxonomic level and enumerated as either total counts or, 

if highly abundant, extrapolated by sub-sampling. A digital still camera was used during 

2003 and 2004 and subsequent analysis revealed that images did not resolve some 

important taxa as well as 35 mm film. Therefore, these years were excluded from this 

analysis. Analyses presented here include data from 1998, 2001, 2002, and 2005. 
5

 

In order to reduce bias from stations where additional transects were performed in a 

particular year, only photographs obtained from the first three transects were used for 

analysis. Photographs containing greater than 20% shadow, where it was not possible to 

identify organisms, were excluded. Imagery obtained during 1998 and 2001 was 

analyzed by a previous observer (Douglas McNaught, Brown University) while we 

analyzed photographs obtained during the 2002 and 2005 sampling years. In order to 

minimize bias in the data produced by two sequential observers we selected and analyzed 

photographs from 1998 and 2001 to comport species identifications. This exercise 

resulted in aggregating some species into higher taxonomic groupings (see Appendix 1, 

Table 2 for a list of taxa used in this study). 

 

Analysis 

 

Multivariate approaches utilized the PRIMER version 5.2.9 software and were used to 

compare community composition and patterns of diversity between habitats, years, and 

closure status (Clarke and Warwick 2001). Similarity matrices, based on the Bray-Curtis 

similarity coefficient, were produced using mean taxon counts per transect for each year. 

Mean abundance of each taxon per transect was used to account for occasional 

differences in transect sample size (e.g., early termination of transects due to weather or 

other logistical constraints). Similarity matrices were produced for gravel and boulder 

habitats separately as well as a global similarity matrix that included both habitat types. 

Non-metric multidimensional scaling (MDS) was used to visualize and evaluate the 

similarity in species composition between treatments. The relative similarity of the 

community structure for each sample site (i.e., each transect, each year at both gravel and 

boulder stations) was plotted in a non-metric space to evaluate the relative similarity of 

each sample location to each other sample site. Analysis of similarities (ANOSIM) tests 

were performed on similarity matrices for each habitat type to evaluate differences in 

contributions of each taxon between habitat type, years sampled and closure status. For 

each ANOSIM test where a significant difference (p<0.05) was identified between 

treatments, similarity percentage - species contributions (SIMPER) analyses were 

performed to identify the top 10 taxa most responsible for the dissimilarity between the 

groups. Standardized un-transformed means for each taxon were used for all SIMPER 

analyses. 

Analysis of variance (ANOVA) using the general linear model was performed to 

elucidate patterns at the level of individual taxa. A global model including three main 

effects (habitat type, year, and closure status) and four interactions terms was used to test 

all taxa for significant differences. Taxa identified in the SIMPER analyses were also 

separated by habitat type where the model included two main effects (year and closure 

status) and one interaction term to test for significant differences. Additional ANOVAs 

were performed using the same global model on structural groups. Species abundance 

data were aggregated a priori into three structural groups in order to generalize the role 

of invertebrate species as structural components of fish habitat: 1) encrusting forms (e.g. 

Didemnum species, encrusting sponges and bryozoans), 2) erect-emergent forms (e.g. 

Terebratulina septentrionalis, Tubularia indivisa, sabellid worms, cerianthid anemones), 
6

and 3) mobile species (e.g., crustaceans, echinoderms). 

 

ANOVAs conducted for individual taxa and structural groups were performed using 

square-root transformed mean abundance per transect data to satisfy homogeneity of 

variance (HOV) requirements. Homogeneity of variance was determined for each taxon 

using either Bartlett’s or Levene’s test statistic depending on violation of the normality 

assumption. As the ANOVA is fairly robust to violations of normality, when the 

normality assumption was not met, Levene’s test statistic for HOV was used (Dytham 

1999, Sokal and Rohlf 2001). 

 

Species diversity measures (i.e., species richness, Shannon-Weiner, and Simpson) were 

calculated for each transect for comparison by univariate ANOVA, again using the 

general linear model. The Shannon-Weiner diversity index responds to changes in the 

number of rare species in a sample while the Simpson diversity index is sensitive to the 

number and abundance of dominant species. Diversity indices were compared based on 

habitat, year, and closure status. Shannon-Weiner and Simpson indices were transformed 

to satisfy HOV requirements (Shannon index was cubed, Simpson index was squared). 

 

K-dominance curves were produced using mean abundance values for each taxa. All 

taxa were ranked by abundance and the percentage of each taxon contributing to the the 

total number of individuals in each year was plotted cumulatively against taxon rank. 

Abundance data was plotted for individual habitat types by year and closure status, for all 

years by closure status, and for closure status by year in order to evaluate changes in the 

relative dominance of the most abundant taxa over time and between habitats within and 

outside the WGOMC. 

Results 

A total of 928 photographic images were analyzed in which 41,690 individuals or 

colonies were counted and assigned to 78 taxonomic categories (species and species 

groups) and 3 structural groups (Appendix 1, Table 2). Based on the structural group 

classification, 11 (14.1%) taxa were encrusting, 39 (50.0%) erect-emergent, and 28 

(35.9%) were mobile. 

The results of two-dimensional non-metric MDS revealed differences in community 

structure based on habitat type, year and closure status (Figure 2). Separate MDS 

procedures for each habitat type by year and closure status illustrate the shifts in 

community structure through time and by closure status (Figure 3a and b). Both gravel 

and boulder stations outside and inside the closure area changed over time from relatively 

similar community structures at the time of closure in 1998. Both the gravel inside and 

outside stations changed through time, but the outside station exhibited less change from 

the initial community state based on relative distances in the MDS plot. The boulder 

stations exhibited the same general type of pattern as the gravel stations. 

A two-way ANOSIM revealed significant differences in community structure between 

years in gravel habitats (R=0.897, p=0.001) and between inside and outside stations 
7

(R=0.880, p=0.001). There were also significant differences in pairwise comparisons of 

all years (p=0.01 for all comparisons; Table 1). Similar results were obtained for boulder 

habitats. There were significant differences between years (R=0.730, p=0.001) and 

between stations inside versus outside the closed area (R=0.759, p=0.001). There were 

also significant differences in invertebrate community composition for all years based on 

pairwise comparisons (p=0.01 for all comparisons; Table 1). A one-way ANOSIM 

comparing taxon composition between gravel and boulder habitats revealed significant 

differences in taxon abundances (R=0.421, p=0.01). 

Of the top ten taxa identified by SIMPER, analysis comparing community composition 

and abundance of individual taxa between gravel and boulder habitats, four taxa were 

more abundant at gravel stations while six taxa were more abundant at boulder stations 

(Table 2). Seven of the 10 taxa contributing to nearly 67% of the dissimilarity between 

boulder and gravel habitats were structure forming invertebrates classified as erectemergent. Analysis of the community composition of gravel habitats based on closure 

status identified a total of four taxa in greater abundance inside the closure and six were 

more abundant outside (Table 3). Here, seven of the taxa contributing to the dissimilarity 

were erect-emergent forms. However, while the erect Iophon spp. and erect bryozoa 

were most abundant inside the closed area Terebratulina septentrionalis, serpulid spp., 

and Cerianthus borealis were most abundant outside. Pairwise comparisons of both 

gravel stations between years using SIMPER revealed differences in taxon abundance 

patterns that did not produce clear directionality in terms of increases in dominant taxa 

(Table 4). 

Similarly, of the top ten taxa contributing to the dissimilarity between boulder stations 

inside and outside the closure area identified by SIMPER analysis three taxa were more 

abundant at stations inside while seven taxa were more abundant outside the closed area 

(Table 5). Seven taxa, contributing to nearly 79% of the dissimilarity between stations, 

were erect-emergent forms. The ascidian Molgula spp. was more than twice as abundant 

at the inside station while the brachiopod Terebratulina septentrionalis was more than six 

times as abundant at the outside station. Two species or forms of Iophon were also more 

abundant outside. Pairwise comparisons of both boulder stations between years using 

SIMPER most strikingly reveal this pattern (Table 6). 

K-dominance curves of gravel communities did not reveal any clear shifts in the patterns 

of dominance of taxa by year or closure status (Figures 4 a, b). However, patterns in the 

curves of boulder communities illustrate a marked increase in the cumulative percent 

dominance of the highest ranked taxa beginning in 2002 at stations inside the closure area 

(Figure 4c). The shape of the dominance curves at the boulder stations outside the 

WGOMC display a slight but consistent increase in abundance of top ranked taxa over 

time (Figure 4d). 

The global ANOVA revealed a total of 20 taxa that differed significantly (p<0.05) in 

abundance based on closure status (Table 7a). Overall, 68 of 79 taxa exhibited one or 

more statistically significant differences based on comparisons of habitat type, year, and 

closure status. Of these a total of 21 taxa were identified to contribute to the 
8

dissimilarity between stations by the previous SIMPER analyses. ANOVAs for each 

habitat type revealed all but five of the 21 taxa had significant differences in abundance 

based on closure status (Table 7b). Changes over time in the mean abundance for the 21 

taxa at each station are summarized in Figure 5. Pairwise comparisons by closure status 

and year for each habitat type identified significant changes in the mean abundance of 20 

taxa (Table 8). Pairwise comparisons also revealed differences in taxon abundance 

patterns that did not produce clear directionality in terms of increases or decreases in 

abundance over time based on closure status at gravel stations. At boulder stations, pairwise comparisons revealed significantly higher abundances of T. septentrionalis and 

associatied encrusting species P. sulfurous at stations outside the closure and significantly 

higher abundances of Asteroidea species at outside stations beginning in 2002 (see 

Appendix 2 for pairwise comparisons across all years). 

Results of the global ANOVA based on structural group designations demonstrated that 

all groups differed significantly in abundance based on habitat type, year, and the habitat 

type by year interaction term (Table 7c). At gravel stations, all structural groups differed 

significantly in abundance by year but only the erect-emergent fauna differed 

significantly (p=0.062) based on closure status (Table 7d). Additionally, both the erectemergent structural group and mobile fauna differed significantly in the year by closure 

status interaction term (Table 7d). At boulder stations, all structural groups differed 

significantly across years and only encrusting fauna differed significantly based on 

closure status (Table 7e). Pairwise comparisons by closure status and across years for 

each habitat type also revealed differences in the abundance of structural groups that did 

not produce clear trends over time (Table 9 and Appendix 2). 

In general, species diversity differed significantly based on habitat types, years, and 

closure status (Figures 6 and 7). The global analysis of the Shannon-Wiener index 

revealed significant differences by year, habitat, and all interaction terms, including 

interaction terms with closure status (ANOVA, all p<0.05), but not closure status alone 

(Table 10). Separate analyses based on habitat type revealed a significant difference in 

the year by closure status interaction term at boulder stations but not at gravel stations. 

Finally, there were significant differences by year for both habitat types. 

The global analysis of the Simpson diversity index revealed significant differences 

(ANOVA, p<0.05) in species diversity for all main effects and interaction terms except 

habitat type (Table 10). Separate analyses based on habitat type revealed significant 

differences (p < 0.01) in diversity across years at gravel habitats, but there were no 

significant differences in closure status, or the interaction of year and closure status 

(Table 10). For boulder habitats, there were significant differences (p < 0.01) by year, 

closure status, and the interaction between year and closure status (Table 10). For each 

habitat type, both the Shannon-Wiener and Simpson index values fluctuated over time 

and were higher at stations located outside the WGOMC with the exception of the 

boulder stations in 2001 (Table 9). At gravel stations, species richness was not 

significantly different based on closure status or the interaction term of year by closure 

status, but was significantly different across years (ANOVA, p < 0.05; Table 10). 

Species richness at gravel stations varied over time, but exhibited an overall decline from 
9

1998 to 2005. At the boulder stations, species richness differed significantly based on the 

year by closure status interaction term (ANOVA, p< 0.05), but there was no significant 

difference between years or closure status (Table 10). Species richness was relatively 

stable across years and among stations, but in 2001 there was a decrease in richness at the 

boulder station outside the closure area (Figure 7c). 

` 

Figure 2. MDS of gravel and boulder habitat community composition. Non-metric multidimensional 

scaling (MDS) ordination for community composition at gravel and boulder stations (based on a 

Bray-Curtis similarity matrix). The naming convention of the stations is 98GIT1, where the first two 

numbers indicate the year, the first letter indicates the habitat type, the second the closure status, 

and the third letter and last number indicate the transect number. The letter X denotes the location 

of the station on the plot. The codes are as follows: G = gravel, B = boulder, I = inside the closed 

area, and O = outside the closed area. The grouping of each substrate type by closure status is 

encircled by solid lines. 
10

Figure 3. Non-metric multidimensional scaling plot illustrating community dynamics by habitat type 

over time; a. gravel habitats, b. boulder habitats. Refer to Figure 2 for label code. The grouping of 

stations by closure status and sampling years is encircled by solid lines. 

a. 

b. 
11

Table 1. Analysis of similarities (ANOSIM) between habitat types; and within each habitat type 

between closure status and sample years. Significant p-values (p<0.100) indicate the analyzed terms 

are signigicantly different from eachother. 

Gravel Boulder

 1998 2001 2002 2005 1998 2001 2002 2005 

1998 p = 0.01 p = 0.01 p = 0.01 1998 p = 0.01 p = 0.01 p = 0.01 

2001 p = 0.01 p = 0.01 2001 p = 0.01 p = 0.01 

2002 p = 0.01 2002 p = 0.01 

2005 2005 

 Closure Status p = 0.001 Closure Status p = 0.001 

 Habitat Type p= 0.001 

Table 2. SIMPER analysis of data by habitat type. Taxa contributing to the dissimilarity (D), based 

on mean abundance (M.A.), between gravel and boulder habitats (average dissimilarity of the mean 

abundance of taxa = 70.49). 

Species Gravel 

M.A. 

Boulder 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 0.56 25.72 12.21 1.24 17.33 17.33 

Erect Iophon spp. 2.36 2.14 6.39 1.07 9.06 26.39 

Terebratulina septentrionalis 1.72 8.40 5.10 1.31 7.24 33.63 

Serpulid spp. 1.89 1.56 3.90 0.83 5.54 39.16 

Cerianthus borealis 1.35 0.68 3.73 1.04 5.29 44.45 

Erect bryozoan spp. 1.33 5.46 3.47 1.36 4.93 49.38 

Encrusting red sponge spp. 2.02 3.93 3.19 1.19 4.52 53.90 

Iophon pattersoni 0.93 4.98 3.13 1.08 4.45 58.35 

Encrusting Iophon spp. 1.67 3.20 3.03 1.25 4.30 62.65 

Encrusting yellow sponge spp. 1.12 0.82 2.77 0.64 3.93 66.57 

Table 3. SIMPER analysis of data for gravel habitats by closure status. Taxa contributing to the 

dissimilarity (D), based on mean abundance (M.A.), at gravel habitats inside and outside the 

WGOMC (average dissimilarity of the mean abundance of taxa = 72.69). 

Taxa Inside 

M.A. 

Outside 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Erect Iophon spp. 2.63 2.08 7.12 1.18 9.80 9.80 

Terebratulina septentrionalis 0.09 3.35 6.53 1.35 8.98 18.78 

Serpulid spp. 1.37 2.41 5.37 0.98 7.39 26.17 

Erect bryozoan spp. 2.58 0.08 5.06 1.70 6.96 33.13 

Cerianthus borealis 0.80 1.90 4.92 1.57 6.76 39.89 

Encrusting red sponge spp. 2.47 1.56 4.36 1.21 6.00 45.89 

Encrusting yellow sponge spp. 1.08 1.16 4.14 0.81 5.70 51.59 

Encrusting Iophon spp. 1.53 1.81 3.79 1.20 5.21 56.81 

Iophon pattersoni 0.01 1.84 3.55 1.22 4.88 61.68 

Holothurian spp. 0.56 0.50 2.71 0.86 3.72 65.41 
12

Table 4. SIMPER analysis of data for gravel stations between years. Taxa contributing to the 

dissimilarity (D), based on mean abundance (M.A.), between years at gravel habitats. 

a. Gravel Years 1998 and 2001 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 65.73)

Taxa 1998 

M.A. 

2001 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Erect Iophon spp. 1.96 4.82 8.96 1.74 13.63 13.63 

Cerianthus borealis 2.55 0.58 8.25 3.82 12.55 26.18 

Holothurian spp. 1.78 0.00 6.71 4.88 10.21 36.39 

Terebratulina septentrionalis 0.76 3.48 6.41 1.10 9.75 46.14 

Asterias vulgaris 0.98 0.30 3.95 1.11 6.00 52.14 

Iophon pattersoni 0.31 1.81 3.37 1.00 5.13 57.27 

Filograna implexa 0.03 1.03 3.24 1.16 4.93 62.21 

Erect bryozoan spp. 0.38 0.85 2.93 1.18 4.45 66.66 

Molgula spp. 0.01 0.91 2.69 1.14 4.09 70.75 

Encrusting bryozoan spp. (pink) 0.66 0.41 2.57 1.10 3.92 74.67 

Encrusting red sponge spp. 0.23 0.19 0.84 1.22 1.28 

b. Gravel Years 1998 and 2002 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 76.22) 

Taxa 1998 

M.A. 

2002 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Encrusting red sponge spp. 0.23 6.38 8.55 3.13 11.21 11.21 

Cerianthus borealis 2.55 1.12 8.14 5.45 10.68 21.89 

Encrusting Iophon spp. 0.06 4.34 6.92 2.66 9.08 30.97 

Holothurian spp. 1.78 0.31 6.24 4.38 8.19 39.16 

Serpulid spp. 0.00 3.23 5.01 5.02 6.57 45.72 

Erect Iophon spp. 1.96 2.23 4.59 1.76 6.02 51.75 

Erect bryozoan spp. 0.38 3.48 4.38 1.13 5.75 57.49 

Asterias vulgaris 0.98 0.03 4.04 0.95 5.30 62.79 

Terebratulina septentrionalis 0.76 1.98 3.62 1.18 4.74 67.54 

Encrusting bryozoan spp. (pink) 0.66 0.07 2.50 0.89 3.29 70.82 

c. Gravel Years 1998 and 2005 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 80.84) 

Taxa 1998 

M.A. 

2005 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Serpulid spp. 0.00 4.32 10.63 1.79 13.15 13.15 

Encrusting yellow sponge spp. 0.00 3.13 8.69 1.47 10.75 23.90 

Cerianthus borealis 2.55 1.14 7.56 2.75 9.35 33.25 

Holothurian spp. 1.78 0.02 6.68 4.85 8.26 41.51 

Erect Iophon spp. 1.96 0.43 6.67 2.44 8.25 49.76 

Encrusting Iophon spp. 0.06 1.88 4.62 1.52 5.71 55.47 

Asterias vulgaris 0.98 0.03 4.05 0.95 5.01 60.48 

Caridean shrimp 0.28 0.43 3.14 0.64 3.89 64.36 

Pagurus spp. 0.04 0.10 2.70 0.50 3.34 67.70 

Encrusting bryozoan spp. (pink) 0.66 0.00 2.56 0.90 3.17 70.87 
13

Table 4. Continued. 

d. Gravel Years 2001 and 2002 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 72.49) 

Taxa 2001 

M.A. 

2002 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Erect Iophon spp. 4.82 2.23 12.29 1.97 16.95 16.95 

Encrusting red sponge spp. 0.19 6.38 9.19 3.48 12.68 29.63 

Terebratulina septentrionalis 3.48 1.98 6.59 1.14 9.09 38.72 

Encrusting Iophon spp. 0.39 4.34 6.19 2.27 8.54 47.27 

Serpulid spp. 0.02 3.23 4.96 4.96 6.85 54.11 

Erect bryozoan spp. 0.85 3.48 4.49 1.21 6.19 60.31 

Iophon pattersoni 1.81 1.31 3.66 1.11 5.06 65.36 

Filograna implexa 1.03 0.42 2.91 1.08 4.02 69.38 

Encrusting yellow sponge spp. 0.02 1.33 2.06 2.32 2.85 72.23 

Molgula spp. 0.91 1.18 1.95 1.04 2.69 74.92 

e. Gravel Years 2001 and 2005 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 82.59) 

Taxa 2001 

M.A. 

2005 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Erect Iophon spp. 4.82 0.43 14.54 2.33 17.61 17.61 

Serpulid spp. 0.02 4.32 10.59 1.78 12.82 30.43 

Encrusting yellow sponge spp. 0.02 3.13 8.66 1.47 10.49 40.91 

Terebratulina septentrionalis 3.48 0.65 6.38 1.00 7.72 48.63 

Encrusting Iophon spp. 0.39 1.88 3.98 1.32 4.82 53.46 

Caridean shrimp 0.03 0.43 3.66 0.71 4.43 57.88 

Iophon pattersoni 1.81 0.28 3.35 0.94 4.06 61.95 

Filograna implexa 1.03 0.39 3.02 1.27 3.66 65.61 

Erect bryozoan spp. 0.85 0.63 2.91 1.24 3.53 69.14 

Encrusting red sponge spp. 0.19 1.27 2.80 1.66 3.39 72.52 

f. Gravel Years 2002 and 2005 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 58.96) 

Taxa 2002 

M.A. 

2005 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Encrusting yellow sponge spp. 1.33 3.13 7.29 1.43 12.36 12.36 

Encrusting red sponge spp. 6.38 1.27 6.58 2.13 11.15 23.51 

Serpulid spp. 3.23 4.32 5.99 1.06 10.16 33.67 

Erect bryozoan spp. 3.48 0.63 4.35 1.18 7.38 41.05 

Encrusting Iophon spp. 4.34 1.88 3.76 1.43 6.37 47.43 

Caridean shrimp 0.45 0.43 3.33 0.67 5.64 53.07 

Terebratulina septentrionalis 1.98 0.65 3.28 1.04 5.56 58.63 

Pagurus spp. 

Cerianthus borealis 

0.00 0.10 2.67 0.48 4.52 63.15 

1.12 1.14 2.50 1.57 4.24 67.39 

Erect Iophon spp. 2.23 0.43 2.31 1.47 3.92 71.31 
14

Table 5. SIMPER analysis of data for boulder habitats based on closure status. Taxa contributing to 

the dissimilarity (D), based on mean abundance (M.A.), at boulder habitats inside and outside the 

WGOMC (average dissimilarity of the mean abundance of taxa = 51.75). 

Taxa Inside 

M.A. 

Outside 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 36.40 15.05 11.44 1.08 22.10 22.10 

Terebratulina septentrionalis 2.25 14.56 6.44 2.17 12.44 34.54 

Iophon pattersoni 0.42 9.54 4.94 1.91 9.56 44.09 

Erect Iophon spp. 1.53 2.75 3.81 0.89 7.37 51.46 

Myxicola infundibulum 2.10 1.27 3.21 1.02 6.21 57.67 

Erect bryozoan spp. 5.00 5.93 2.99 1.34 5.78 63.45 

Ophiopholis aculeata 1.55 2.16 2.65 0.88 5.13 68.58 

Pseudosuberites sulfureus 1.70 4.59 1.94 1.59 3.75 72.33 

Encrusting Iophon spp. 2.71 3.69 1.65 1.18 3.19 75.52 

Asteroidea unidentifiable 1.95 1.18 1.54 1.35 2.97 78.49 
15

Table 6. SIMPER analysis of data for boulder stations between years. Taxa contributing to the 

dissimilarity (D), based on mean abundance (M.A.), between years at boulder stations. 

a. Boulder Years 1998 and 2001 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 45.92)

Taxa 1998 

M.A. 

2001 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Erect Iophon spp. 1.86 2.82 5.60 1.08 12.18 12.18 

Ophiopholis aculeata 6.76 0.52 4.79 1.69 10.43 22.61 

Myxicola infundibulum 2.63 2.39 4.42 1.21 9.62 32.23 

Erect bryozoan spp. 7.07 1.41 3.91 1.61 8.52 40.75 

Terebratulina septentrionalis 7.26 1.89 3.28 1.82 7.13 47.88 

Molgula spp. 9.01 2.94 3.27 1.13 7.13 55.01 

Asteroidea unidentifiable 1.36 1.57 2.35 4.38 5.11 60.12 

Pseudosuberites sulfureus 3.60 0.91 1.92 1.39 4.19 64.30 

Iophon pattersoni 2.40 0.70 1.84 1.37 4.01 68.32 

Encrusting Iophon spp. 1.84 1.10 1.47 1.44 3.19 71.51 

b. Boulder Years 1998 and 2002 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 49.26) 

Taxa 1998 

M.A. 

2002 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 9.01 19.13 8.41 1.69 17.07 17.07 

Ophiopholis aculeata 6.76 0.01 5.99 2.30 12.17 29.24 

Terebratulina septentrionalis 7.26 8.11 4.69 1.56 9.53 38.77 

Erect bryozoan spp. 7.07 2.83 4.15 1.65 8.42 47.18 

Iophon pattersoni 2.40 5.93 3.83 1.39 7.77 54.95 

Erect Iophon spp. 1.86 3.28 2.58 1.15 5.23 60.18 

Encrusting Iophon spp. 1.84 4.69 2.27 1.16 4.62 64.80 

Myxicola infundibulum 2.63 1.29 2.22 1.14 4.50 69.30 

Pseudosuberites sulfureus 

Filograna implexa 

3.60 2.06 2.07 1.35 4.20 73.51 

0.46 2.08 1.52 1.12 3.09 76.60 

c. Boulder Years 1998 and 2005 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 53.12)

Taxa 1998 

M.A. 

2005 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 9.01 71.82 15.38 1.21 28.96 28.96 

Ophiopholis aculeata 6.76 0.13 5.93 2.28 11.17 40.13 

Terebratulina septentrionalis 7.26 16.36 5.15 1.64 9.70 49.83 

Iophon pattersoni 2.40 10.90 3.75 1.39 7.06 56.88 

Erect bryozoan spp. 7.07 10.54 3.27 1.35 6.16 63.05 

Myxicola infundibulum 2.63 0.43 2.35 1.00 4.42 67.47 

Pseudosuberites sulfureus 3.60 6.01 2.13 1.35 4.01 71.48 

Erect Iophon spp. 1.86 0.61 1.46 0.89 2.75 74.23 

Polymastia hispida 1.71 0.00 1.43 1.06 2.70 76.93 

Serpulid spp. 0.09 3.93 1.39 1.90 2.62 79.55 
16

Table 6. Continued. 

d. Boulder Years 2001 and 2002 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 48.25) 

Taxa 2001 

M.A. 

2002 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 2.94 19.13 9.52 1.68 19.72 19.72 

Erect Iophon spp. 2.82 3.28 5.59 1.24 11.59 31.32 

Terebratulina septentrionalis 1.89 8.11 4.54 2.20 9.41 40.72 

Myxicola infundibulum 2.39 1.29 4.42 1.04 9.16 49.88 

Iophon pattersoni 0.70 5.93 3.86 1.33 7.99 57.87 

Erect bryozoan spp. 1.41 2.83 2.37 1.26 4.90 62.78 

Encrusting Iophon spp. 1.10 4.69 1.89 1.13 3.92 66.70 

Asteroidea unidentifiable 1.57 2.32 1.60 1.55 3.32 70.02 

Filograna implexa 0.74 2.08 1.46 1.39 3.02 73.04 

Pseudosuberites sulfureus 0.91 2.06 1.28 1.42 2.65 75.69 

e. Boulder Years 2001 and 2005 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 58.87) 

Taxa 2001 

M.A. 

2005 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 2.94 71.82 16.69 1.29 28.36 28.36 

Erect Iophon spp. 2.82 0.61 5.83 0.99 9.91 38.27 

Terebratulina septentrionalis 1.89 16.36 5.14 2.86 8.73 47.00 

Myxicola infundibulum 2.39 0.43 5.13 1.18 8.72 55.72 

Iophon pattersoni 0.70 10.90 3.78 1.36 6.41 62.13 

Asteroidea unidentifiable 1.57 1.01 3.19 6.38 5.41 67.54 

Erect bryozoan spp. 1.41 10.54 2.64 1.43 4.49 72.03 

Pseudosuberites sulfureus 0.91 6.01 1.66 1.54 2.81 74.85 

Encrusting red sponge spp. 1.25 7.46 1.41 1.31 2.40 77.25 

Encrusting Iophon spp. 1.10 5.17 1.38 1.46 2.34 79.58 

f. Boulder Years 2002 and 2005 Dissimilarity (Average dissimilarity of the mean 

abundance of taxa: 41.05) 

Taxa 2002 

M.A. 

2005 

M.A. 

Mean 

D 

St. 

Dev.

Contributing

% 

Cumulative

% 

Molgula spp. 19.13 71.82 12.70 1.28 30.93 30.93 

Terebratulina septentrionalis 8.11 16.36 5.24 1.15 12.77 43.70 

Iophon pattersoni 5.93 10.90 4.03 1.15 9.83 53.52 

Erect Iophon spp. 3.28 0.61 2.76 1.09 6.73 60.25 

Erect bryozoan spp. 2.83 10.54 2.23 1.48 5.42 65.68 

Encrusting Iophon spp. 4.69 5.17 2.02 1.15 4.93 70.61 

Pseudosuberites sulfureus 2.06 6.01 1.61 1.47 3.91 74.52 

Asteroidea unidentifiable 2.32 1.01 1.60 1.62 3.89 78.41 

Filograna implexa 2.08 1.67 1.49 1.20 3.62 82.03 

Myxicola infundibulum 1.29 0.43 1.04 0.97 2.53 84.56 
17

Figure 4. K dominance curves for gravel and boulder habitats by year and closure status. Taxon 

rank plotted against the cumulative dominance based on percent representation for gravel (a and b) 

and boulder (c and d) stations located inside (a and c) and outside (b and d) the WGOMC by year. 

Legend : = 1998, = 2001, = 2002, and = 2005. 

a. 

b. 

c. 

d. 

Species Rank 

Cumuative Dominance (%) 
18

Table 7. Analysis of Variance for individual taxa and structural groups. Results of ANOVAs for: a. 

all taxa; b. 21 taxa identified by SIMPER analysis; c. structural groups (global model); d. structural 

groups in gravel habitats; and e. structural groups in boulder habitats. P-values are given for the 

terms that are significant (p<0.10); n.s. = p > 0.10. The terms are: habitat type (H), year (Y), and 

closure status (CS). 

a. 

Taxa H Y CS H*Y H*CS Y*CS H*Y* 

CS 

Aplysilla spp. 0.009 n.s. n.s. n.s. n.s. n.s. n.s. 

Encrusting yellow sponge spp. n.s. <0.001 n.s. n.s. n.s. n.s. n.s. 

Haliclona oculata n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Haliclona urceola <0.001 0.019 0.035 n.s. 0.012 0.038 n.s. 

Leuconia spp. n.s. <0.001 0.001 n.s. n.s. <0.001 n.s. 

Hymedesmia sp. 1 n.s. 0.010 n.s. n.s. n.s. n.s. n.s. 

Hymedesmia sp. 2 n.s. 0.014 n.s. n.s. n.s. n.s. 0.017 

Encrusting red sponge spp. <0.001 <0.001 n.s. <0.001 n.s. 0.007 n.s. 

Encrusting Iophon spp. <0.001 <0.001 n.s. 0.033 n.s. n.s. n.s. 

Erect Iophon spp. n.s. <0.001 n.s. n.s. n.s. n.s. n.s. 

Iophon pattersoni <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 

Pseudosuberites sulfurous <0.001 0.037 <0.001 <0.001 n.s. 0.006 0.015 

Isodictya palmate n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Mycale spp. n.s. <0.001 n.s. n.s. n.s. 0.017 <0.001 

Suberites spp. n.s. <0.001 n.s. n.s. n.s. n.s. n.s. 

Plocambionida ambigua 0.005 0.008 n.s. 0.001 n.s. <0.001 <0.001 

Polymastia hispida 0.007 0.001 n.s. 0.001 n.s. n.s. n.s. 

Polymastia spp. <0.001 n.s. 0.029 0.014 n.s. n.s. n.s. 

Sycon ciliate n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Phakellia ventilabrum n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Alcyonium digitatum n.s. <0.001 n.s. n.s. n.s. n.s. n.s. 

Closed Anemone 0.042 0.010 n.s. 0.010 n.s. n.s. n.s. 

Bolocera tuediae n.s. n.s. n.s. n.s. n.s. 0.007 0.008 

Cerianthus borealis 0.001 <0.001 <0.001 0.012 <0.001 <0.001 n.s. 

Cerianthiopsis americanus n.s. <0.001 n.s. n.s. 0.038 0.022 n.s. 

Tubularia indivisa n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Urticina feline 0.028 n.s. n.s. n.s. n.s. n.s. n.s. 

Urticina spp. n.s. <0.001 n.s. n.s. n.s. n.s. n.s. 

Erect bryozoan spp. <0.001 0.001 0.001 <0.001 0.001 <0.001 n.s. 

Tubilopora sp. 1 <0.001 n.s. n.s. n.s. n.s. n.s. n.s. 

Encrusting Bryozoan spp. pink <0.001 <0.001 <0.001 <0.001 0.002 0.007 0.024 

Encrusting Bryozoan spp. white 0.004 <0.001 n.s. n.s. n.s. n.s. n.s. 

Terebratulina septentrionalis <0.001 0.033 <0.001 <0.001 0.028 0.002 <0.001 

Amphiporus angulatus n.s. 0.025 n.s. n.s. n.s. n.s. n.s. 

Nereis sp. 0.054 0.016 n.s. 0.016 n.s. 0.016 0.016 

Filograna implexa <0.001 0.003 <0.001 0.012 n.s. n.s. n.s. 

Myxicola infundibulum <0.001 0.001 n.s. n.s. <0.001 0.007 0.001 

Serpulid spp. n.s. <0.001 n.s. 0.002 n.s. <0.001 0.011 

Sabelid spp. white 0.027 <0.001 n.s. n.s. n.s. n.s. n.s. 

Sabelid spp. red 0.035 n.s. 0.014 n.s. n.s. n.s. n.s. 

Calliostoma spp. n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Colus spp. n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Epitonium spp. 0.005 n.s. n.s. n.s. n.s. n.s. n.s. 

Neptunea lyrata decemcostata n.s. n.s. 0.008 0.038 n.s. n.s. n.s. 
19

Table 7. Continued. 

a. Continued. 

Taxa H Y CS H*Y H*CS Y*CS H*Y* 

CS

Astarte spp. n.s. 0.004 n.s. 0.025 n.s. n.s. n.s. 

Cyclocardia borealis n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Modiolus modiolus 0.002 

n.s. 

0.032 n.s. n.s. 0.029 

0.022 

n.s. 

n.s. 

n.s. 

Nudibranch 0.001 0.022 n.s. n.s. 

Hyas spp. n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Pagurus spp. <0.001 0.044 <0.001 0.013 0.030 n.s. 0.015 

Caridean Shrimp 0.023 <0.001 n.s. <0.001 0.014 n.s. n.s. 

Balanus balanus 0.001 n.s. 0.026 n.s. n.s. n.s. 0.032 

Asterias vulgaris 0.046 <0.001 0.002 <0.001 <0.001 <0.001 0.002 

Crossaster papposus 0.002 n.s. n.s. 0.036 0.010 n.s. n.s. 

Henricia sanguinolenta <0.001 <0.001 n.s. 0.003 n.s. n.s. n.s. 

Hippasteria phrygiana n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Porania insignis <0.001 n.s. n.s. n.s. n.s. n.s. n.s. 

Solaster endeca n.s. 0.025 0.043 0.022 n.s. n.s. n.s. 

Stephanasterias albula 0.054 0.016 n.s. 0.016 n.s. n.s. n.s. 

Asteroidea unidentifiable <0.001 n.s. n.s. 0.006 <0.001 0.019 n.s. 

Pteraster militaria n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Ophiopholis aculeate <0.001 <0.001 n.s. <0.001 n.s. n.s. n.s. 

Echinarachnius parma n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Hathrometra spp. 0.008 <0.001 n.s. <0.001 n.s. n.s. n.s. 

Cucumaria frondosa n.s. n.s. n.s. 0.010 n.s. n.s. n.s. 

Psolus spp. n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Holothurian spp. <0.001 <0.001 n.s. 0.001 n.s. n.s. n.s. 

Strongylocentrotus 

droebachiensis n.s. n.s. n.s. n.s. n.s. n.s. n.s. 

Ascidia callosa n.s. <0.001 n.s. n.s. n.s. n.s. n.s. 

Boltenia echinata 0.066 <0.001 n.s. 0.023 n.s. n.s. 0.043 

Boltenia ovifera 0.004 n.s. n.s. n.s. 0.025 n.s. n.s. 

Ciona intestinalis <0.001 n.s. 0.007 0.004 n.s. n.s. 0.023 

Didemnum albidum n.s. n.s. n.s. 0.013 n.s. n.s. n.s. 

Didemnum sp.1 <0.001 <0.001 0.002 0.009 n.s. 0.021 n.s. 

Halocynthia pyriformis 0.037 0.002 n.s. n.s. 0.005 n.s. n.s. 

Trididemnum solidum 0.035 0.082 n.s. n.s. n.s. 0.023 0.023 

Molugula spp. <0.001 <0.001 0.021 <0.001 0.030 n.s. n.s. 

Synocium pulmonaria n.s. 0.001 n.s. n.s. n.s. n.s. n.s. 
20

Table 7. Continued. 

b.

Taxa Identified by SIMPER 

Gravel Boulder 

Y CS Y*CS Y CS Y*CS 

Encrusting yellow sponge <0.001 n.s. n.s. <0.001 n.s. n.s. 

Encrusting red sponge <0.001 n.s. 0.008 <0.001 0.04 n.s. 

Encrusting Iophon spp. <0.001 n.s. n.s. <0.001 n.s. n.s. 

Iophon pattersoni n.s. <0.001 n.s. <0.001 <0.001 <0.001 

Pseudosuberites sulfureus n.s. <0.001 n.s. 0.004 0.002 0.009 

Erect Iophon spp. <0.001 n.s. 0.013 n.s. n.s. n.s. 

Polymastia hispida NA NA NA 0.003 n.s. n.s. 

Cerianthus borealis <0.001 <0.001 0.002 0.001 n.s. 0.056 

Erect bryozan spp. 0.01 <0.001 0.006 <0.001 n.s. 0.009 

Encrusting bryozoan spp. (pink) n.s. 0.016 n.s. 0.035 0.073 0.035 

Terebratulina septentrionalis 0.088 <0.001 n.s. <0.001 <0.001 <0.001 

Filograna implexa 0.002 0.001 n.s. 0.041 0.044 0.092 

Myxicola infundibulum 0.029 <0.001 0.02 0.026 0.046 0.005 

Serpulid spp. <0.001 0.032 <0.001 <0.001 n.s. 0.055 

Pagurus spp. 0.008 0.001 0.017 n.s. n.s. n.s. 

Caridean shrimp <0.001 0.003 0.011 0.004 n.s. n.s. 

Asterias vulgaris <0.001 <0.001 <0.001 0.039 n.s. n.s. 

Asteroidea unidentifiable <0.001 <0.001 <0.001 <0.001 <0.001 0.009 

Ophiopholis aculeata 0.001 n.s. 0.025 <0.001 n.s. n.s. 

Holothurian spp. <0.001 n.s. n.s. <0.001 n.s. n.s. 

Molgula spp. <0.001 n.s. n.s. <0.001 0.03 n.s. 

c.

Structural Group H Y CS H*Y H*CS Y*CS H*Y* 

CS 

Encrusting <0.001 <0.001 n.s. 0.001 n.s. n.s. n.s. 

Erect-emergent <0.001 <0.001 n.s. <0.001 n.s. n.s. n.s. 

Mobile <0.001 <0.001 n.s. <0.001 n.s. n.s. 0.015 

d. 

Gravel 

Structural Group Y CS Y*CS 

 Encrusting <0.001 n.s. n.s. 

Erect-emergent 0.012 0.062 0.094 

Mobile <0.001 n.s. 0.023 

e. 

Boulder 

Structural Group Y CS Y*CS 

Encrusting <0.001 0.064 n.s. 

Erect-emergent <0.001 n.s. n.s. 

Mobile <0.001 n.s. n.s. 
21

Figure 5. Mean abundance of selected taxa over time based on habitat type and closure status. The 

mean abundance of the 21 taxa identified by SIMPER analysis are plotted by year and habitat type. 

Stations inside the WGOMC are plotted as solid lines and outside stations are plotted as dotted lines. 

Year 

Mean Abundance 

Gravel

Encrusting yellow sponge spp.

0.00

1.00

2.00

3.00

4.00

1998 2001 2002 2005

Gravel 

Encrusting red sponge spp. 

0.00

2.00

4.00

6.00

8.00

10.00

1998 2001 2002 2005

Gravel 

Encrusting Iophon spp. 

0.00

2.00

4.00

6.00

1998 2001 2002 2005

Boulder

Encrusting yellow sponge spp. 

0.00

0.50

1.00

1.50

2.00

2.50

1998 2001 2002 2005

Boulder

Encrusting red sponge spp. 

0.00

2.00

4.00

6.00

8.00

10.00

1998 2001 2002 2005

Boulder

Encrusting Iophon spp.

0.00

2.00

4.00

6.00

8.00

1998 2001 2002 2005
22

Figure 5. Continued. Year 

Mean Abundance 

Gravel 

Cerianthus borealis

0.00

1.00

2.00

3.00

1998 2001 2002 2005

Gravel 

Erect bryozan spp.

0.00

2.00

4.00

6.00

8.00

1998 2001 2002 2005

Boulder 

Cerianthus borealis

0.00

0.50

1.00

1.50

2.00

1998 2001 2002 2005

Boulder

Erect bryozan spp.

0.00

5.00

10.00

15.00

20.00

1998 2001 2002 2005

Gravel 

Polymastia hispida

0.00

0.50

1.00

1998 2001 2002 2005

Boulder 

Polymastia hispida 

0.00

0.50

1.00

1.50

2.00

2.50

1998 2001 2002 2005

Gravel 

Iophon pattersoni

0.00

1.00

2.00

3.00

4.00

1998 2001 2002 2005

Gravel 

Pseudosuberites sulfureus

0.00

0.50

1.00

1.50

1998 2001 2002 2005

Gravel 

Erect Iophon spp.

0.00

2.00

4.00

6.00

8.00

1998 2001 2002 2005

Boulder

Iophon pattersoni

0.00

10.00

20.00

30.00

1998 2001 2002 2005

Boulder 

Pseudosuberites sulfureus 

0.00

2.00

4.00

6.00

8.00

10.00

12.00

1998 2001 2002 2005

Boulder 

Erect Iophon spp.

0.00

1.00

2.00

3.00

4.00

5.00

1998 2001 2002 2005
23

Figure 5. Continued. 

Gravel 

Myxicola infundibulum 

0.00

0.50

1.00

1.50

1998 2001 2002 2005

Boulder 

Myxicola infundibulum 

0.00

1.00

2.00

3.00

4.00

5.00

1998 2001 2002 2005

Gravel 

Pagurus spp.

0.00

0.05

0.10

0.15

0.20

0.25

1998 2001 2002 2005

Boulder 

Pagurus spp.

0.00

0.01

0.01

0.02

0.02

1998 2001 2002 2005

Gravel 

Serpulid spp.

0.00

2.00

4.00

6.00

8.00

1998 2001 2002 2005

Boulder 

Serpulid spp. 

0.00

1.00

2.00

3.00

4.00

5.00

6.00

1998 2001 2002 2005

Gravel 

Encrusting bryozoan spp. (pink) 

0.00

0.50

1.00

1.50

1998 2001 2002 2005

Gravel 

Terebratulina septentrionalis 

0.00

2.00

4.00

6.00

8.00

1998 2001 2002 2005

Boulder 

Terebratulina septentrionalis

0.00

10.00

20.00

30.00

40.00

1998 2001 2002 2005

Boulder

Encrusting bryozoan spp. (pink)

0.00

0.02

0.04

0.06

1998 2001 2002 2005

Gravel 

Filograna implexa

0.00

0.50

1.00

1.50

2.00

1998 2001 2002 2005

Boulder 

Filograna implexa

0.00

1.00

2.00

3.00

4.00

1998 2001 2002 2005

Year 

Mean Abundance 
24

Figure 5. Continued. 

Gravel 

Ophiopholis aculeata

0.00

0.50

1.00

1.50

1998 2001 2002 2005

Gravel 

Holothurian spp.

0.00

0.50

1.00

1.50

2.00

1998 2001 2002 2005

Gravel 

Molgula spp.

0.00

0.50

1.00

1.50

1998 2001 2002 2005

Boulder 

Ophiopholis aculeata

0.00

2.00

4.00

6.00

8.00

10.00

1998 2001 2002 2005

Boulder

Holothurian spp.

0.00

0.20

0.40

0.60

0.80

1998 2001 2002 2005

Boulder 

Molgula spp. 

0.00

20.00

40.00

60.00

80.00

100.00

120.00

1998 2001 2002 2005

Gravel 

Caridean shrimp 

0.00

0.20

0.40

0.60

0.80

1998 2001 2002 2005

Gravel 

Asterias vulgaris

0.00

0.50

1.00

1.50

2.00

2.50

1998 2001 2002 2005

Boulder 

Caridean shrimp

0.00

0.20

0.40

0.60

0.80

1998 2001 2002 2005

Boulder

Asterias vulgaris

0.00

0.05

0.10

0.15

0.20

0.25

1998 2001 2002 2005

Gravel 

Asteroidea unidentifiable

0.00

0.10

0.20

0.30

0.40

0.50

1998 2001 2002 2005

Boulder

Asteroidea unidentifiable

0.00

1.00

2.00

3.00

4.00

1998 2001 2002 2005

Year 

Mean Abundance 
25

Table 8. Pairwise comparisons for taxonomic categories in gravel and boulder habitat. Cell values 

indicate the percent of the 21 taxa identified by SIMPER analysis. For comparisons between years (a 

thru d), the first value represents the percent of taxa that had higher abundance in the earlier year. 

The second value represents the percent of taxa that had higher abundance in the later year. For 

comparisons based on closure status (e and d), the first value represents the percent of taxa that had 

higher abundance inside the closed area. The second value represents the percent of taxa that had 

higher abundance outside the closed area. 

a. Gravel In 

 1998 2001 2002 2005 

1998 

2001 14%, 19% 

2002 19%, 33% 19%, 33% 

2005 24%, 5% 19%, 5% 14%, 0% 

c. Gravel Out 

 1998 2001 2002 2005 

1998 

2001 5%, 0% 

2002 5%, 24% 0%, 24% 

2005 14%, 14% 0%, 24% 19%, 10% 

e. Gravel In versus Gravel Out 

 1998 2001 2002 2005 

1998 14%, 14% 

2001 19%, 5% 

2002 19%, 14% 

2005 0%, 24% 

b. Boulder In 

 1998 2001 2002 2005 

1998 

2001 33%, 10% 

2002 14%, 14% 5%, 19% 

2005 19%, 14% 5%, 14% 5%, 0% 

d. Boulder Out 

 1998 2001 2002 2005 

1998 

2001 24%, 0% 

2002 10%, 24% 5%, 38% 

2005 10%, 38% 5%, 43% 5%, 24% 

f. Boulder In versus Boulder Out 

 1998 2001 2002 2005 

1998 5%, 5% 

2001 5%, 5% 

2002 5%, 14% 

2005 5%, 24% 
26 

Table 9. Pairwise comparisons for structural group categories in gravel and boulder habitat. Cell 

values indicate the percent of the 3 structural groups exhibiting significant differences between years. 

For comparisons between years (a thru d), the first value represents the percent of taxa that had 

higher abundance in the earlier year. The second value represents the percent of taxa that had 

higher abundance in the later year. For comparisons based on closure status (e and d), the first value 

represents the percent of taxa that had higher abundance inside the closed area. The second value 

represents the percent of taxa that had higher abundance outside the closed area. 

a. Gravel In 

1998 2001 2002 2005 

1998 

2001 33%, 33% 

2002 33%, 66% 0%, 66% 

2005 0%, 33% 0%, 0% 33%, 0% 

c. Gravel Out 

 1998 2001 2002 2005 

1998 

2001 0%, 0% 

2002 0%, 33% 0%, 33% 

2005 33%, 33% 33%, 33% 33%, 0% 

e. Gravel In versus Gravel Out 

 1998 2001 2002 2005 

1998 0%, 0% 

2001 0%, 33% 

2002 33%, 0% 

2005 0%, 33% 

b. Boulder In 

1998 2001 2002 2005 

1998 

2001 66%, 0% 

2002 33%, 0% 0%, 100% 

2005 33%, 66% 0%, 66% 0%, 66% 

d. Boulder Out 

 1998 2001 2002 2005 

1998 

2001 100%, 0% 

2002 33%, 33% 0%, 66% 

2005 33%, 66% 33%, 66% 0%, 100% 

f. Boulder In versus Boulder Out 

 1998 2001 2002 2005 

1998 0% , 0% 

2001 0%, 0% 

2002 33%, 33% 

2005 33%, 0% 
30 

Figure 6. Species diversity indices for gravel habitats by year. a. Shannon-Weinner Index; b. 

Simpson Index; c. Species Richness. Legend: = inside, = outside. 

Year 

c. 

b. 

a. 

Species Diversity Index 
31 

Figure 7. Species diversity indices for boulder habitats by year. a. Shannon-Weinner Index; b. 

Simpson Index; c. Species Richness. Legend: = inside, = outside. 

c. 

b. 

a. 

Year 

Species Diversity Index 
32 

Table 10. Analysis of variance for species diversity indices. Results of ANOVAs for: a. global 

model; and b. gravel and boulder habitats separately. P-values are given for the terms that are 

significant (p<0.10); n.s. = p > 0.10. The terms are: habitat type (H), year (Y), and closure status 

(CS). 

a. 

Diversity Index H Y CS H*Y H*CS Y*CS H*Y* 

CS 

Shannon-Wiener 0.006 < 0.001 n.s < 0.001 n.s 0.014 0.006 

Simpson n.s < 0.001 0.001 < 0.001 0.048 0.003 < 0.001

Species Richness < 0.001 0.004 n.s 0.032 n.s n.s 0.058 

b. 

 Gravel Boulder 

Diversity Index Y CS Y*CS Y CS Y*CS 

Shannon-Wiener 0.002 n.s n.s < 0.001 n.s 0.003 

Simpson < 0.001 n.s n.s < 0.001 0.007 0.002 

Richness 0.017 n.s n.s n.s n.s 0.021 
33 

Discussion 

Overall these results demonstrate that community structure over the seven years since 

closure has been dynamic across both habitat types, as well as within and outside the 

WGOMC, despite a high degree of similarity between paired habitat stations at the time 

of closure. Multivariate and univariate comparisons of community structure, populations 

of component taxa, and patterns of diversity between habitat types inside and outside the 

closed area across years all demonstrate a response to the closure, but not in ways that are 

normally predicted from previous closed area studies. 

Despite hard substratum resources in both boulder and gravel habitats, community 

structure was different between habitat types across all years. The effects of variations 

in: 1) flow regime, 2) growth and competition of suspension feeders, and 3) shelter 

resources from predators can contribute to such differences (Denny 1988, Lesser et al. 

1995). We infer that boulder habitats have a much higher degree of variation in flow 

fields over hard substratum due to wide variation in size and density of boulders whereas 

gravel habitats have lower flow field variation because they are generally planar and with 

cobble-pebble distributed as a flat pavement (based on interpretation of maps; Valentine 

et al. 2003). 

Community structure changed across time both inside and outside the WGOMC 

suggesting recovery without resilience at both habitat types (Paine et al. 1998, Gunderson 

2000). In gravel and boulder habitats, the community structure changed significantly 

across the time series at sites outside the WGOMC indicating the communities are not 

resilient to disturbance effects (Palumbi et al. 2008, Levin and Lubchenco 2008). The 

significant differences between gravel stations inside and outside the closure based on 

multiple analyses suggest recovery is occurring inside the closure but, to date, there is no 

clearly definable stable community structure. In contrast, at boulder stations there was a 

clear shift in community composition inside the closure area over time that we infer was 

due to local processes (i.e., competitive interactions). 

Most taxa at the gravel station inside the closed area did not exhibit clear trajectories in 

abundance, and both increased and decreased over the time series. Only caridean shrimp 

exhibited a significant overall increase in abundance inside the WGOMC (see Appendix 

2 for details on individual taxa). In contrast, encrusting red sponges, Molgula spp., 

Suberites spp., and Serpulid spp. significantly increased in abundance at boulder stations 

inside the closed area. It is difficult to ascribe drivers to the patterns of abundance for 

most taxa. However, the significantly greater abundance of T. septentrionalis at boulder 

stations outside the closed area and the distinct shift in dominance to Molgula spp. inside 

the closed area may be attributed to competitive interactions. Brachiopod species 

worldwide have exhibited declining trends in abundance when productivity of local 

ecosystems increased. Due to the slower metabolisms and filtration rates of brachiopods, 

modern suspension-feeders (i.e., mollusks, asidians) gain a competitive advantage where 

local processes drive species interactions (Thayer, 1986; Bambach 1993; Rhodes and 

Thompson, 1993; McKinney and Hageman, 2006; Clapham and Bottjer 2007). Here we 

conclude that disturbance by fishing outside the closed area is the dominant driver of the 
34 

community structure allowing brachiopods to dominate space resources, whereas inside 

the closed area competitive interactions are favoring Molgula spp. and other modern 

suspension-feeders reducing the dominant role of T. septentrionalis. Interestingly, there 

were no clear shifts in dominance or clear directionality in the abundance of particular 

taxa at gravel stations, although pairwise comparisons revealed T. septentrionalis was 

significantly higher in abundance outside the closed area in 2005. The pattern with T. 

septentrionalis suggests local processes are driving the changes in the epifaunal 

community structure between stations inside and outside the closure area. Terebratulina 

septentrionalis may be a good indicator species for chronic disturbance in the western 

Gulf of Maine. 

Community structure within the closed area has yet to reach any stable composition. 

Further, community structure outside the closed area has not remained in what might be 

considered a consistent impacted state. While community composition tended to be more 

similar within each station than between each year, the pattern of similarity from 2005 

data suggest a greater degree of difference in composition between replicates from gravel 

and boulder stations inside the closed area than paired stations outside the closed area. 

This pattern also suggests the community-level processes, such as predation and 

competition, may be driving species composition inside the closed area (i.e., contributing 

to greater variation in species distributions within stations). This is in contrast to broader 

spatial scale disturbance processes, produced either by natural events or by fishing 

activities that dominate at stations outside the closed area. 

This study is predicated on the fact that the human disturbance regime (largely in the 

form of bottom trawl and sink gillnet fishing gear) has greatly diminished inside the 

closure while continuing at a higher level immediately outside the closure. However, it is 

important to note that the fishing pressure outside the closed area has not been constant 

over time and has varied with changes in both the regulatory regime and fishing 

economics (Murawski et al. 2005). For example, the outside gravel station at the time of 

closure was impacted primarily by trawl gear. Due to the implementation of seasonal 

closures and other fishing sector interactions, the site is now primarily impacted by fixed 

gillnets. Further, over the course of time the level of overall fishing effort has decreased 

regionally as well as locally (Murawski et al. 2005). The effect of such variation in 

fishing effort and gear types on the dynamics of community structure reported here is 

unknown. 

Interestingly, structural guilds and population trajectories of component taxa changed 

over time in unpredictable ways. We predicted that erect-emergent (structure forming) 

invertebrates would increase in abundance over time within the protected area due to 

elimination of fishing gear disturbance. At the gravel stations the erect-emergent fauna 

was significantly different based on closure status, but abundance was generally greater 

at the outside station over the time series (with the exception of 2002). Whereas there 

was no significant difference in the erect-emergent fauna based on closure status at 

boulder stations over time, there was a significant difference in the abundance of 

encrusting forms, with greater numbers found outside the closure area (with the exception 

of 2001). No significant differences were identified in mobile fauna based on closure 
35 

status for either habitat type. Perhaps most importantly, at the initiation of the time series 

the sites outside the closure area of both habitats had greater abundances of the erectemergent and encrusting structural groups and this pattern remained essentially 

unchanged over time. 

The expected increase in species diversity at stations within the WGOMC was not 

observed by the end of the study at either the gravel or boulder stations. The Kdominance curves for gravel habitat illustrate little difference in patterns of dominance 

over the seven years. In contrast to the results of prior studies (Collie et al. 1997, Collie 

et al. 2005), the gravel stations outside the WGOMC had higher diversity than the gravel 

stations inside, although this relationship varied between years. Initially species diversity 

decreased from 1998 to 2001, then increased from 2001 to 2002, and then decreased 

again from 2002 to 2005. The same patterns where observed in species richness. The 

difference in diversity over years may be due either to competitive interactions between 

species within each station or the recruitment of new species which require habitats with 

reduced levels of disturbance. 

Species diversity also differed significantly at boulder stations across years and by 

closure status. At the station inside the closure area diversity increased from 1998 to 

2001 and subsequently decreased through 2005 when calculated using Shannon-Wiener’s 

diversity index. This increase likely indicates an increase in abundance of rare species 

(e.g., not detected in earlier surveys) or an initial recruitment event post-closure. The 

subsequent decline in diversity indicates recruitment leveled off and competitive 

interactions or predation, or both, dominated thereby decreasing the abundance of rare 

species. Except for a slight increase in 2001, diversity declined inside the closure area 

across the time series based upon Simpson’s diversity index. This decline reflects the 

observed changes in patterns of dominance. The K-dominance curves illustrate these 

changes in species diversity over time. The marked upward shift of the K-dominance 

curve at the inside station over time is due to the large increase in abundance of Molgula 

spp.. This shift in the contribution of dominant species would result in a reduction in 

evenness of the community and reduce diversity as calculated by Simpson’s index. At 

the station outside the closure area, there was a continual, minor decline in diversity when 

calculated by both Shannon-Wiener and Simpson’s diversity indices. There was also a 

marked decrease in species richness in 2001. The observed declines in species diversity 

outside the closure area are likely linked to the decrease in species richness in 2001 

which subsequently increased to near the 1998 level in 2002 and 2005. 

These observations neither support nor reject the assumption that cessation or reduction 

of fishing will allow populations and communities to recover to a climax community 

state. As of 2005, a successional end point of the communities at gravel and boulder 

stations remains unclear. These findings do indicate that the WGOMC is having a 

significant impact on invertebrate community structure, and that the community inside 

the closure area on both boulder and gravel habitats is recovering from chronic fishing 

gear impacts. The lack of stability suggests that community structure under protected and 

impacted regimes is dynamic and that “recovery” of the seafloor community has not, and 

perhaps will not, reach a stable climax state. At this time it is not possible to match the 
36 

dynamics of communities at SBNMS to either of the two general models of community 

dynamics. Neither gravel nor boulder stations inside the closed area have reached a 

stable community state. Only the results of continued monitoring will enable us to define 

the type of community model that operates in this region of the Gulf of Maine. 

While identifying the type of community model driving the changes we have documented 

in the community structure remains elusive, the changes we have identified indicate the 

WGOMC is conserving biodiversity in both gravel and boulder habitats. Because the 

essential elements of Essential Fish Habitat have yet to be defined for all stages of 

managed fish species (e.g., Lindolm et al 1999, Gotceitas and Brown 1993), the 

conservation of biodiversity through the use of MPAs across a broad spectrum of habitats 

remains a valuable management tool (Alcala et al 2005, Hart 2006, Auster 2001). 

Further studies are necessary to evaluate which attributes of invertebrate community 

structure are important to maintain and enhance the productivity of managed fish species 

across a wide variety of habitat types. 

Acknowledgements 

This work was funded primarily through a contract from Perot Systems Government 

Services with funds from NOAA’s National Marine Sanctuary Program. Overall the 

Seafloor Habitat Recovery Monitoring Program benefited from funding and support from 

multiple organizations throughout this seven year time series and these include the 

National Undersea Research Center at the University of Connecticut, U.S. Geological 

Survey, National Marine Fisheries Service Northeast Fisheries Science Center, 

University of Maine, University of Massachusetts, and Brown University. We thank the 

masters and crews of the following vessels for outstanding support in the field: FV 

Christopher Andrew, NOAA Ship Ferrel, NOAA Ship Nancy Foster, FV Isabel S, and 

RV Connecticut. We also thank, in particular, the following individuals: Dennis Arbige, 

Dan Blackwood, Craig Bussel, Paul Donaldson, Betsy Grannis, Les Kaufman, Chiu-Yen 

Kuo, Douglas McNaught, Page Valentine, Les Watling, and Jon Witman for their 

collaborations and help during the course of this larger project. We express our particular 

gratitude to Brad Barr and Michael Sissenwine who had the vision to make limited 

funding resources available for the first of these cruises and to Craig MacDonald, the 

current SBNMS Superintendent, for continued support. The reviews by Drs. J. Evan 

Ward and Eric Schultz greatly improved the manuscript. The opinions expressed herein 

are those of the authors and do not necessarily reflect the opinions NOAA or any of the 

other agencies or institutions that participated in or funded this study. 

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Appendix 1: Sampling and Data Collection Details 

Table 1. Sampling cruise schedule from 1998 through 2005. Unless otherwise noted 

gravel and boulder habitats were sampled on the same cruise. 

Year Sampling Begin Sampling End 

1998 July 5 July 8 

2001 August 7 August 13 

2002 April 30 (gravel), June 6 (boulder) May 1 (gravel), June 6 (boulder) 

2003 June 18 June 20 

2004 July 17 July 20 

2005 June 11 June 13 

Table 2. Taxonomic categories (species and species groups*) and structural groups 

identified in image analysis. 

Phylum Taxon Groups Structural 

Group Comment 

Porifera Aplysilla sp. Encrusting

 Encrusting yellow sponge spp. Encrusting Halichondria sp. 1 + Iophon yellow 

encrusting + Leptosia yellow 

 Haliclona oculata Erect-emergent 

 Haliclona urceola Erect-emergent 

 Leuconia spp. Erect-emergent 

 Hymedesmia sp. 1 Encrusting blue 

 Hymedesmia sp. 2 Encrusting orange 

 Encrusting red sponge spp. Encrusting Hymedesmia sp. 3 (red) + Myxilla 

fimbriata + Leptosia pink 

 Encrusting Iophon spp. Encrusting Iophon white encrusting + Porifera sp. 1 

and sp. 2 + Porifera gray 

 Erect Iophon spp. Erect-emergent 

 Iophon pattersoni Erect-emergent 

 Pseudosuberites sulfurous Erect-emergent 

 Isodictya palmate Erect-emergent 

 Mycale spp. Erect-emergent Mycale lingua + Mycale placoides + 

Porifer yellow smooth 

 Suberites spp. Erect-emergent Mycale lobata + Suberites spp. 

 Plocamionida ambigua Erect-emergent 

 Polymastia hispida Erect-emergent 

 Polymastia spp. Erect-emergent Polymastia infrapilosa + Polymastia sp. 1 

 Sycon ciliate Erect-emergent 

 Phakellia ventilabrum Erect-emergent 

 Alcyonium digitatum Erect-emergent 

Cnidaria Closed anemone Erect-emergent 

 Bolocera tuediae Erect-emergent 

 Cerianthus borealis Erect-emergent 

 Ceriantheoposis americanus Erect-emergent 

 Tubularia indivisa Erect-emergent 

 Urticina feline Erect-emergent 

 Urticina spp. Erect-emergent 
42 

Table 2. Continued. 

Phylum Taxon Groups Structural 

Group Comment

Bryozoa Erect bryozoan spp. Erectemergent Caberea ellisii + Idmidronea atlantica 

 Tubulipora sp. 1 Erectemergent 

 Encrusting bryozoan spp. Encrusting pink 

 Encrusting bryozoan spp. Encrusting white 

Brachiopoda Terebratulina septentrionalis Erectemergent 

Nermertea Amphiporus angulatus Mobile 

 Nereis sp. Mobile 

Annelida Filograna implexa Erectemergent 

 Myxicola infundibulum Erectemergent 

 Serpulid spp. Erectemergent 

 Sabellid spp. (white) Erectemergent 

 Sabellid spp. (red) Erectemergent 

Mollusca Calliostoma spp. Mobile 

 Colus spp. Mobile 

 Epitonium spp. Mobile 

 Neptunea lyrata decemcostata Mobile 

 Astarte spp. Mobile 

 Cyclocardia borealis Mobile 

 Modiolus modiolus Erectemergent 

 Nudibranch Mobile 

Crustacea Hyas spp. Mobile 

 Pagurus spp. Mobile 

 Caridean shrimp Mobile 

 Balanus balanus Erectemergent 

Echinodermata Asterias vulgaris Mobile 

 Crossaster papposus Mobile 

 Henricia sanguinolenta Mobile 

 Hippasteria phrygiana Mobile 

 Porania insignis Mobile 

 Solaster endeca Mobile 

 Stephanasterias albula Mobile 

 Asteroidea unidentifiable Mobile Asteroidea spp. + Leptasterias spp. 

 Pteraster militaria Mobile 

 Ophiopholis aculeate Mobile 

 Echinarachnius parma Mobile 

 Hathrometra spp. Mobile 

 Cucumaria frondosa Mobile 
43 

Table 2. Continued. 

Phylum Taxon Groups Structural 

Group Comment

Echinodermata Psolus spp. Mobile 

continued Holothurian spp. Mobile 

Strongylocentrotus 

droebachiensis Mobile 

Urochordata Ascidia callosa Erectemergent 

Boltenia echinata Erectemergent 

Boltenia ovifera Erectemergent 

Ciona intestinalis Erectemergent 

Didemnum albidum Encrusting 

Didemnum sp. 1 Encrusting 

Halocynthia pyriformis Erectemergent 

Trididemnum solidum Encrusting 

Molgula spp. 

Erectemergent Molgula spp. + Gray and silt-covered 

individuals* 

Synoicum pulmonaria Erectemergent 

*Silt covered tunicates were counted as individuals for 1998 and 2001 data, and counted 

as blocks for 2002 and 2005 data. The following conversion was used to convert block 

counts to individual counts: Silt-covered individual = individual count; Silt-covered 

blocks low density (1-3 individuals per block) = total number of blocks multiplied by 1; 

Silt-covered blocks medium density (4-10 individuals per block) = total number of blocks 

multiplied by 4; and Silt-covered blocks high density (>10 individuals per block) = total 

number of blocks multiplied by 11. The sum of the block conversions was then entered 

as the total number of individuals for the taxa. 
44 

Appendix 2: Details of Pairwise Comparisons 

Table 1. Results of pairwise comparisons of data for taxonomic categories in gravel 

habitat. a. comparison of 1998 and 2001; b. comparisons of 1998 and 2002; c. 

comparisons of 1998 and 2005; d. comparisons of 2001 and 2002; e. comparisons of 

2001 and 2005; and f. comparisons of 2002 and 2005. 

a. 

SPECIES GI v. GO 2001 GI 1998 v. GI 2001 GO 1998 v. GO 2001 

Encrusting yellow sponge NS NS NS 

Encrusting red sponge NS NS NS 

Encrusting Iophon spp. NS NS NS 

Iophon pattersoni NS NS NS 

Pseudosuberites sulfureus GO=Higher, p=0.051 NS NS 

Erect Iophon spp. NS 2001=Higher, p=0.005 NS 

Polymastia hispida NS NS NS 

Cerianthus borealis GI=Higher, p=0.019 1998=Higher, p=0.001 NS 

Erect bryozan spp. GI=Higher, p=0.021 NS NS 

Encrusting bryozoan (pink) NS NS NS 

Terebratulina septentrionalis NS NS NS 

Filograna implexa NS 2001=Higher, p=0.005 NS 

Myxicola infundibulum NS NS NS 

Serpulid spp. NS NS NS 

Pagurus spp. NS NS NS 

Caridean shrimp NS NS 1998=Higher, p=0.01 

Asterias vulgaris GI=Higher, p=0.005 1998=Higher, p=0.015 NS 

Asteroidea unidentifiable NS NS NS 

Ophiopholis aculeata NS 2001=Higher, p=0.01 NS 

Holothurian tentacles GI=Higher, p=0.002 1998=Higher, p=0.002 NS 

Molgula spp. NS 2001=Higher, p=0.03 NS 
45 

Table 1. Continued. 

b. 

TAXA GI v. GO 2002 GI 1998 v. GI 2002 GO 1998 v. GO 2002 

Encrusting yellow sponge NS 2002=Higher, p<0.001 2002=Higher, p=0.036 

Encrusting red sponge GI=Higher, p=0.016 2002=Higher, p=0.002 2002=Higher, p=0.001 

Encrusting Iophon spp. NS 2002=Higher, p=0.01 2002=Higher, p<0.001 

Iophon pattersoni GO=Higher, p=.115 NS NS 

Pseudosuberites sulfureus GO=Higher, p=0.046 NS NS 

Erect Iophon spp. GI=Higher, p=0.028 2002=Higher, p=0.05 NS 

Polymastia hispida NS NS NS 

Cerianthus borealis NS 1998=Higher, p=0.011 NS 

Erect bryozan spp. GI=Higher, p=0.047 NS NS 

Encrusting bryozoan (pink) NS NS NS 

Terebratulina septentrionalis NS NS NS 

Filograna implexa NS 2002=Higher, p=0.041 NS 

Myxicola infundibulum NS NS 1998=Higher, p=0.043 

Serpulid spp. GI=Higher, p<0.001 2002=Higher, p<0.001 2002=Higher, p<0.001 

Pagurus spp. NS 1998=Higher, p=0.016 NS 

Caridean shrimp GO=Higher, p=0.019 NS 2002=Higher, p=0.035 

Asterias vulgaris NS 1998=Higher, p=0.005 NS 

Asteroidea unidentifiable NS NS NS 

Ophiopholis aculeata NS NS NS 

Holothurian tentacles NS 1998=Higher, p=0.001 NS 

Molgula spp. NS 2002=Higher, p=0.01 NS 

c. 

TAXA GI v. GO 1998 GI v. GO 2005 GI 1998 v. GI 2005 GO 1998 v. GO 2005

Encrusting yellow sponge NS NS NS 2005 Higher, p=0.009

Encrusting red sponge GO Higher, p=0.039 NS NS 2005 Higher, p=0.052

Encrusting Iophon spp. GI Higher, p=0.047 NS NS NS 

Iophon pattersoni NS NS NS NS 

Pseudosuberites sulfureus NS GO Higher, p=0.017 NS NS 

Erect Iophon spp. NS NS 1998 Higher, p=0.057 1998 Higher, p=0.047

Polymastia hispida NS NS NS NS 

Cerianthus borealis NS GO Higher, p=0.008 1998 Higher, p=0.001 NS 

Erect bryozan spp. NS NS NS NS 

Encrusting bryozoan (pink) NS NS 1998 Higher, p=0.055 NS 

Terebratulina septentrionalis NS GO Higher, p=0.005 NS NS 

Filograna implexa GI Higher, p=0.016 NS NS NS 

Myxicola infundibulum GO Higher, p=0.031 NS NS 1998 Higher, p=0.048

Serpulid spp. NS GO Higher, p=0.023 NS 2005 Higher, p=0.004

Pagurus spp. NS NS NS NS 

Caridean shrimp GO Higher, p=0.06 NS 2005 Higher, p=0.009 NS 

Asterias vulgaris GI Higher, p=0.006 NS 1998 Higher, p=0.006 NS 

Asteroidea unidentifiable NS GO Higher, p=0.053 NS NS 

Ophiopholis aculeata NS NS NS NS 

Holothurian tentacles NS NS 1998 Higher, p=0.001 1998 Higher, p=0.058

Molgula spp. NS NS NS NS 
46 

Table 1. Continued. 

d.

TAXA GI 2001 v. GI 2002 GO 2001 v. GO 2002 

Encrusting yellow sponge 2002=Higher, p<0.001 2002=Higher, p=0.039 

Encrusting red sponge 2002=Higher, p=0.002 2002=Higher, p=0.001 

Encrusting Iophon spp. 2002=Higher, p=0.01 2002=Higher, p<0.001 

Iophon pattersoni NS NS 

Pseudosuberites sulfureus NS NS 

Erect Iophon spp. 2002=Higher, p=0.05 NS 

Polymastia hispida NS NS 

Cerianthus borealis 2001=Higher, p=0.011 NS 

Erect bryozan spp. NS NS 

Encrusting bryozoan (pink) NS NS 

Terebratulina septentrionalis NS NS 

Filograna implexa 2002=Higher, p=.041 NS 

Myxicola infundibulum NS NS 

Serpulid spp. 2002=Higher, p<0.001 2002=Higher, p<0.001 

Pagurus spp. 2001=Higher, p=0.016 NS 

Caridean shrimp NS 2002=Higher, p=0.003 

Asterias vulgaris 2001=Higher, p<0.001 NS 

Asteroidea unidentifiable NS NS 

Ophiopholis aculeata NS NS 

Holothurian tentacles 2001=Higher,p=0.011 NS 

Molgula spp. 2002=Higher, p=0.01 NS 

e. 

TAXA GI 2001 v. GI 2005 GO 2001 v. GO 2005 

Encrusting yellow sponge NS 2005=Higher, p=0.009 

Encrusting red sponge NS 2005=Higher, p=0.053 

Encrusting Iophon spp. NS NS 

Iophon pattersoni NS NS 

Pseudosuberites sulfureus NS NS 

Erect Iophon spp. NS NS 

Polymastia hispida NS NS 

Cerianthus borealis 2001=Higher, p=0.001 2005=Higher, p=0.051 

Erect bryozan spp. NS NS 

Encrusting bryozoan (pink) 2001=Higher, p=0.047 NS 

Terebratulina septentrionalis NS NS 

Filograna implexa NS NS 

Myxicola infundibulum NS NS 

Serpulid spp. NS 2005=Higher, p=0.005 

Pagurus spp. NS NS 

Caridean shrimp 2005=Higher, p=0.001 2005=Higher, p=0.001 

Asterias vulgaris 2001=Higher, p=0.001 NS 

Asteroidea unidentifiable NS NS 

Ophiopholis aculeata NS NS 

Holothurian tentacles 2001=Higher, p=0.002 NS 

Molgula spp. NS NS 
47 

Table 1. Continued. 

f. 

TAXA GI 2002 v. GI 2005 GO 2002 v. GO 2005 

Encrusting yellow sponge NS 2005=Higher, p=0.046 

Encrusting red sponge 2002=Higher, p=0.004 2002=Higher, p=0.023 

Encrusting Iophon spp. NS 2002=Higher, p=0.011 

Iophon pattersoni NS NS 

Pseudosuberites sulfurous NS NS 

Erect Iophon spp. 2002=Higher, p=0.007 2002=Higher, p=0.041 

Polymastia hispida NS NS 

Cerianthus borealis NS NS 

Erect bryozan spp. NS NS 

Encrusting bryozoan (pink) NS NS 

Terebratulina septentrionalis NS NS 

Filograna implexa NS NS 

Myxicola infundibulum NS NS 

Serpulid spp. NS 2005=Higher, p=0.017 

Pagurus spp. NS NS 

Caridean shrimp NS 2002=Higher, p=0.032 

Asterias vulgaris NS NS 

Asteroidea unidentifiable NS NS 

Ophiopholis aculeate NS NS 

Holothurian tentacles NS NS 

Molgula spp. 2002=Higher, p=0.01 NS 
48 

Table 2. Results of pairwise comparisons of data for taxonomic categories in boulder 

habitat. a. comparison of 1998 and 2001; b. comparisons of 1998 and 2002; c. 

comparisons of 1998 and 2005; d. comparisons of 2001 and 2002; e. comparisons of 

2001 and 2005; and f. comparisons of 2002 and 2005. 

a. 

TAXA BI v. BO 2001 BI 1998 v. BI 2001 BO 1998 v. BO 2001 

Encrusting yellow sponge NS NS NS 

Encrusting red sponge NS 1998=Higher, p=0.016 1998=Higher, p=0.012 

Encrusting Iophon spp. NS NS NS 

Iophon pattersoni NS NS 1998=Higher, p=0.011 

Pseudosuberites sulfureus NS NS NS 

Erect Iophon spp. NS NS NS 

Polymastia hispida NS NS NS 

Cerianthus borealis BO=Higher, p=0.027 1998=Higher, p=0.043 NS 

Erect bryozan spp. NS 1998=Higher, p=0.011 1998=Higher, p=0.009 

Encrusting bryozoan (pink) BI=Higher, p=0.016 2001=Higher, p=0.016 NS 

Terebratulina septentrionalis NS NS 1998=Higher, p=0.007 

Filograna implexa NS NS NS 

Myxicola infundibulum NS 1998=Higher, p=0.044 NS 

Serpulid spp. NS 2001=Higher, p=0.029 NS 

Pagurus spp. NS NS NS 

Caridean shrimp NS 1998=Higher, p=0.031 NS 

Asterias vulgaris NS NS NS 

Asteroidea unidentifiable NS NS NS 

Ophiopholis aculeata NS 1998=Higher, p<0.001 1998=Higher, p=0.052 

Holothurian tentacles NS 1998=Higher, p=0.045 NS 

Molgula spp. NS NS NS 
49 

Table 2. Continued. 

b. 

TAXA BI v. BO 2002 BI 1998 v. BI 2002 BO 1998 v. BO 2002 

Encrusting yellow sponge NS 2002=Higher, p=0.003 2002=Higher, p=0.007 

Encrusting red sponge NS NS NS 

Encrusting Iophon spp. NS NS 2002=Higher, p=0.013 

Iophon pattersoni BO=Higher, p<0.001 NS 2002=Higher, p=0.001 

Pseudosuberites sulfureus BO=Higher, p=0.028 NS NS 

Erect Iophon spp. NS NS NS 

Polymastia hispida NS NS NS 

Cerianthus borealis NS NS NS 

Erect bryozan spp. NS NS 1998=Higher, p=0.039 

Encrusting bryozoan (pink) NS NS NS 

Terebratulina septentrionalis BO=Higher, p<0.001 NS NS 

Filograna implexa NS 2002=Higher, p=0.04 NS 

Myxicola infundibulum NS NS NS 

Serpulid spp. NS NS 2002=Higher, p=0.001 

Pagurus spp. NS NS NS 

Caridean shrimp NS 1998=Higher, p=0.031 NS 

Asterias vulgaris NS NS 2002=Higher, p=0.035 

Asteroidea unidentifiable BI=Higher, p=0.027 2002=Higher, p=0.01 NS 

Ophiopholis aculeata NS 1998=Higher, p<0.001 1998=Higher, p=0.041 

Holothurian tentacles NS 1998=Higher, p=0.045 NS 

Molgula spp. NS NS NS 

c. 

TAXA BI v. BO 1998 BI v. BO 2005 BI 1998 v. BI 2005 BO 1998 v. BO 2005

Encrusting yellow sponge NS NS NS 2005 Higher, p=0.012

Encrusting red sponge NS BO Higher, p=0.037 2005 Higher, p=0.009 2005 Higher, p=0.003

Encrusting Iophon spp. NS NS NS 2005 Higher, p=0.014

Iophon pattersoni BO Higher, p=0.001 BO Higher, p<0.001 NS 2005 Higher, p<0.001

Pseudosuberites sulfureus NS BO Higher, p=0.006 NS 2005 Higher, p=0.033

Erect Iophon spp. NS NS NS NS 

Polymastia hispida NS NS NS NS 

Cerianthus borealis NS NS 1998 Higher, p=0.041 NS 

Erect bryozan spp. NS BO Higher, p=0.057 NS 2005 Higher, p=0.018

Encrusting bryozoan (pink) NS NS NS NS 

Terebratulina septentrionalis NS BO Higher, p<0.001 NS 2005 Higher, p=0.002

Filograna implexa NS NS NS NS 

Myxicola infundibulum BI Higher, p=0.026 NS 1998 Higher, p=0.025 NS 

Serpulid spp. NS NS 2005 Higher, p=0.009 2005 Higher, p=0.024

Pagurus spp. NS NS NS NS 

Caridean shrimp NS NS NS NS 

Asterias vulgaris NS NS NS NS 

Asteroidea unidentifiable NS BI Higher, p=0.045 NS 1998 Higher, p=0.003

Ophiopholis aculeata NS NS 1998 Higher, p<0.001 1998 Higher, 0.044 

Holothurian tentacles NS NS 1998 Higher, p=0.049 NS 

Molgula spp. NS NS 2005 Higher, p=0.003 NS 
50 

Table 2. Continued. 

d. 

TAXA BI 2001 v. BI 2002 BO 2001 v. BO 2002 

Encrusting yellow sponge 2002=Higher, p=0.005 2002=Higher, p=0.003 

Encrusting red sponge 2002=Higher, p=0.023 2002=Higher, p=0.003 

Encrusting Iophon spp. NS 2002=Higher, p=0.004 

Iophon pattersoni NS 2002=Higher, p<0.001 

Pseudosuberites sulfurous NS 2002=Higher, p=0.029 

Erect Iophon spp. NS NS 

Polymastia hispida NS NS 

Cerianthus borealis NS 2001=Higher, p=0.011 

Erect bryozoan spp. NS NS 

Encrusting bryozoan (pink) 2001=Higher, p=0.016 NS 

Terebratulina septentrionalis NS 2002=Higher, p<0.001 

Filograna implexa 2002=Higher, p=0.051 NS 

Myxicola infundibulum NS NS 

Serpulid spp. NS 2002=Higher, p<0.001 

Pagurus spp. NS NS 

Caridean shrimp NS NS 

Asterias vulgaris NS NS 

Asteroidea unidentifiable 2002=Higher, p=0.023 NS 

Ophiopholis aculeate NS NS 

Holothurian tentacles NS NS 

Molgula spp. NS 2002=Higher, p=0.048 

e. 

TAXA BI 2001 v. BI 2005 BO 2001 v. BO 2005 

Encrusting yellow sponge NS 2005=Higher, p=0.004 

Encrusting red sponge 2005=Higher, p=0.001 2005=Higher, p=0.001 

Encrusting Iophon spp. 2005=Higher, p=0.04 2005=Higher, p=0.003 

Iophon pattersoni NS 2005=Higher, p<0.001 

Pseudosuberites sulfurous NS 2005=Higher, p=0.004 

Erect Iophon spp. NS NS 

Polymastia hispida NS NS 

Cerianthus borealis NS NS 

Erect bryozoan spp. NS 2005=Higher, p=0.002 

Encrusting bryozoan (pink) 2001=Higher, p=0.016 NS 

Terebratulina septentrionalis NS 2005=Higher, p<0.001 

Filograna implexa NS 2005=Higher, p=0.05 

Myxicola infundibulum NS NS 

Serpulid spp. 2005=Higher, p=0.022 2005=Higher, p=0.023 

Pagurus spp. NS NS 

Caridean shrimp NS NS 

Asterias vulgaris NS NS 

Asteroidea unidentifiable NS 2001=Higher, p=0.002 

Ophiopholis aculeate NS NS 

Holothurian tentacles NS NS 

Molgula spp. NS NS 
51 

Table 2. Continued. 

f. 

TAXA BI 2002 v. BI 2005 BO 2002 v. BO 2005 

Encrusting yellow sponge NS NS 

Encrusting red sponge NS 2005=Higher, p=0.005 

Encrusting Iophon spp. NS NS 

Iophon pattersoni NS 2005=Higher, p=0.001 

Pseudosuberites sulfurous NS 2005=Higher, p=0.012 

Erect Iophon spp. NS NS 

Polymastia hispida NS NS 

Cerianthus borealis NS NS 

Erect bryozan spp. NS 2005=Higher, p=0.003 

Encrusting bryozoan (pink) NS NS 

Terebratulina septentrionalis NS 2005=Higher, p=0.002 

Filograna implexa NS NS 

Myxicola infundibulum NS NS 

Serpulid spp. NS NS 

Pagurus spp. NS NS 

Caridean shrimp NS NS 

Asterias vulgaris NS NS 

Asteroidea unidentifiable 2002=Higher, p=0.025 2002=Higher, p=0.024 

Ophiopholis aculeate NS NS 

Holothurian tentacles NS NS 

Molgula spp. NS NS 
52 

Table 3. Results of pairwise comparisons of data for structural groups in gravel habitat. 

a. comparison of 1998 and 2001; b. comparisons of 1998 and 2002; c. comparisons of 

1998 and 2005; d. comparisons of 2001 and 2002; e. comparisons of 2001 and 2005; and 

f. comparisons of 2002 and 2005. 

a.

GROUP GI v. GO 2001 GI 1998 v. GI 2001 GO 1998 v. GO 2001 

Encrusting NS NS NS 

Erect-Emergent NS 2001=Higher, p=0.006 NS 

Mobile GO = Higher, p=0.026 1998=Higher, p=0.001 NS 

b. 

GROUP GI v. GO 2002 GI 1998 v. GI 2002 GO 1998 v. GO 2002 

Encrusting GI = Higher, p=0.058 2002=Higher, p=0.002 2002=Higher, p<0.001 

Erect-Emergent NS 2002=Higher, p=0.007 NS 

Mobile NS 1998=Higher, p=0.004 NS 

c. 

GROUP GI v. GO 1998 GI v. GO 2005 GI 1998 v. GI 2005 GO 1998 v. GO 2005 

Encrusting NS NS NS 2005=Higher, p=0.002 

Erect-Emergent NS GO = Higher, p=0.087 NS NS 

Mobile NS NS 2005=Higher, p=0.010 1998=Higher, p=0.047 

d. 

GROUP GI 2001 v. GI 2002 GO 2001 v. GO 2002 

Encrusting 2002=Higher, p=0.001 2002=Higher, p=0.003 

Erect-Emergent 2002=Higher, p=0.057 NS 

Mobile NS NS 

e. 

GROUP GI 2001 v. GI 2005 GO 2001 v. GO 2005 

Encrusting NS 2005=Higher, p=0.019 

Erect-Emergent NS NS 

Mobile NS 2001=Higher, p=0.008 

f. 

GROUP GI 2002 v. GI 2005 GO 2002 v. GO 2005 

Encrusting NS 2002=Higher, p=0.042 

Erect-Emergent 2002=Higher, p=0.045 NS 

Mobile NS NS 
53 

Table 4. Results of pairwise comparisons of data for structural groups in boulder habitat. 

a. comparison of 1998 and 2001; b. comparisons of 1998 and 2002; c. comparisons of 

1998 and 2005; d. comparisons of 2001 and 2002; e. comparisons of 2001 and 2005; and 

f. comparisons of 2002 and 2005. 

a.

GROUP BI v. BO 2001 BI 1998 v. BI 2001 BO 1998 v. BO 2001 

Encrusting NS NS 1998 = Higher, p=0.011 

Erect-Emergent NS 1998 = Higher, p=0.002 1998 = Higher, p=0.006 

Mobile NS 1998 = Higher, p<0.001 1998 = Higher, p=0.013 

b. 

GROUP BI v. BO 2002 BI 1998 v. BI 2002 BO 1998 v. BO 2002 

Encrusting BO=Higher, p=0.069 NS 2002=Higher, p=0.033 

Erect-Emergent NS NS NS 

Mobile BI=Higher, p=0.030 1998=Higher, p=0.001 1998=Higher, p=0.011 

c. 

GROUP BI v. BO 1998 BI v. BO 2005 BI 1998 v. BI 2005 BO 1998 v. BO 2005 

Encrusting NS NS 2005 = Higher, p=0.049 2005 = Higher, p=0.002 

Erect-Emergent NS NS 2005 = Higher, p=0.095 2005 = Higher, p=0.001 

Mobile NS BI=Higher, p=0.041 1998 = Higher, p<0.001 1998=Higher, p=0.006 

 

d. 

GROUP BI 2001 v. BI 2002 BO 2001 v. BO 2002 

Encrusting 2002=Higher, p=0.006 2002=Higher, p=0.001 

Erect-Emergent 2002=Higher, p=0.004 2002=Higher, p=0.004 

Mobile 2002=Higher, p=0.072 NS 

e. 

GROUP BI 2001 v. BI 2005 BO 2001 v. BO 2005 

Encrusting 2005=Higher, p=0.006 2005=Higher, F=751.19, p<0.001 

Erect-Emergent 2005=Higher, p=0.033 2005=Higher, F=132.61, p<0.001 

Mobile NS 2001=Higher, F=7.52, p=0.052 

f. 

GROUP BI 2002 v. BI 2005 BO 2002 v. BO 2005 

Encrusting 2005=Higher, p=0.094 2005=Higher, p=0.029 

Erect-Emergent NS 2005=Higher, p=0.006 

Mobile 2002=Higher, Fp=0.032 2002=Higher, p=0.028 

 
1 

ONMS CONSERVATION SERIES PUBLICATIONS 

To date, the following reports have been published in the Marine Sanctuaries Conservation 

Series. All publications are available on the Office of National Marine Sanctuaries Web site 

(http://www.sanctuaries.noaa.gov/). 

Stellwagen Bank Marine Historical Ecology Final Report (ONMS-10-04) 

Variation in Planning-Unit Size and Patterns of Fish Diversity: Implications for Design of 

Marine Protected Areas (ONMS-10-03) 

Examples of Ecosystem-Based Management in National Marine Sanctuaries: Moving from 

Theory to Practice (ONMS-10-02) 

The Application Of Observing System Data In California Current Ecosystem Assessments 

(ONMS-10-01) 

Reconciling Ecosystem-Based Management and Focal Resource Conservation in the 

Papahānaumokuākea Marine National Monument (ONMS-09-04) 

Preliminary Comparison of Natural Versus Model-predicted Recovery of Vessel-generated 

Seagrass Injuries in Florida Keys National Marine Sanctuary (ONMS-09-03) 

A Comparison of Seafloor Habitats and Associated Benthic Fauna in Areas Open and Closed 

to Bottom Trawling Along the Central California Continental Shelf (ONMS-09-02) 

Chemical Contaminants, Pathogen Exposure and General Health Status of Live and BeachCast Washington Sea Otters (Enhydra lutris kenyoni) (ONMS-09-01) 

Caribbean Connectivity: Implications for Marine Protected Area Management (ONMS-08-

07) 

Knowledge, Attitudes and Perceptions of Management Strategies and Regulations of 

FKNMS by Commercial Fishers, Dive Operators, and Environmental Group Members: A 

Baseline Characterization and 10-year Comparison (ONMS-08-06) 

First Biennial Ocean Climate Summit: Finding Solutions for San Francisco Bay Area's Coast 

and Ocean (ONMS-08-05) 

A Scientific Forum on the Gulf of Mexico: The Islands in the Stream Concept (NMSP-08-04) 

M/V ELPIS Coral Reef Restoration Monitoring Report Monitoring Events 2004-2007 Florida 

Keys National Marine Sanctuary Monroe County, Florida (NMSP-08-03) 

CONNECTIVITY Science, People and Policy in the Florida Keys National Marine Sanctuary 

(NMSP-08-02) 
2 

M/V ALEC OWEN MAITLAND Coral Reef Restoration Monitoring Report Monitoring 

Events 2004-2007 Florida Keys National Marine Sanctuary Monroe County, Florida (NMSP08-01) 

Automated, objective texture segmentation of multibeam echosounder data - Seafloor survey 

and substrate maps from James Island to Ozette Lake, Washington Outer Coast. (NMSP-07-

05) 

Observations of Deep Coral and Sponge Assemblages in Olympic Coast National Marine 

Sanctuary, Washington (NMSP-07-04) 

A Bioregional Classification of the Continental Shelf of Northeastern North America for 

Conservation Analysis and Planning Based on Representation (NMSP-07-03) 

M/V WELLWOOD Coral Reef Restoration Monitoring Report Monitoring Events 2004-2006 

Florida Keys National Marine Sanctuary Monroe County, Florida (NMSP-07-02) 

Survey report of NOAA Ship McArthur II cruises AR-04-04, AR-05-05 and AR-06-03: 

Habitat classification of side scan sonar imagery in support of deep-sea coral/sponge 

explorations at the Olympic Coast National Marine Sanctuary (NMSP-07-01) 

2002 - 03 Florida Keys National Marine Sanctuary Science Report: An Ecosystem Report 

Card After Five Years of Marine Zoning (NMSP-06-12) 

Habitat Mapping Effort at the Olympic Coast National Marine Sanctuary - Current Status and 

Future Needs (NMSP-06-11) 

M/V CONNECTED Coral Reef Restoration Monitoring Report Monitoring Events 2004-

2005 Florida Keys National Marine Sanctuary Monroe County, Florida (NMSP-06-010) 

M/V JACQUELYN L Coral Reef Restoration Monitoring Report Monitoring Events 2004-

2005 Florida Keys National Marine Sanctuary Monroe County, Florida (NMSP-06-09) 

M/V WAVE WALKER Coral Reef Restoration Baseline Monitoring Report - 2004 Florida 

Keys National Marine Sanctuary Monroe County, Florida (NMSP-06-08) 

Olympic Coast National Marine Sanctuary Habitat Mapping: Survey report and classification 

of side scan sonar data from surveys HMPR-114-2004-02 and HMPR-116-2005-01 (NMSP06-07) 

A Pilot Study of Hogfish (Lachnolaimus maximus Walbaum 1792) Movement in the Conch 

Reef Research Only Area (Northern Florida Keys) (NMSP-06-06) 

Comments on Hydrographic and Topographic LIDAR Acquisition and Merging with 

Multibeam Sounding Data Acquired in the Olympic Coast National Marine Sanctuary 

(ONMS-06-05) 

Conservation Science in NOAA's National Marine Sanctuaries: Description and Recent 

Accomplishments (ONMS-06-04) 
3 

Normalization and characterization of multibeam backscatter: Koitlah Point to Point of the 

Arches, Olympic Coast National Marine Sanctuary - Survey HMPR-115-2004-03 (ONMS06-03) 

Developing Alternatives for Optimal Representation of Seafloor Habitats and Associated 

Communities in Stellwagen Bank National Marine Sanctuary (ONMS-06-02) 

Benthic Habitat Mapping in the Olympic Coast National Marine Sanctuary (ONMS-06-01) 

Channel Islands Deep Water Monitoring Plan Development Workshop Report (ONMS-05-

05) 

Movement of yellowtail snapper (Ocyurus chrysurus Block 1790) and black grouper 

(Mycteroperca bonaci Poey 1860) in the northern Florida Keys National Marine Sanctuary as 

determined by acoustic telemetry (MSD-05-4) 

The Impacts of Coastal Protection Structures in California's Monterey Bay National Marine 

Sanctuary (MSD-05-3) 

An annotated bibliography of diet studies of fish of the southeast United States and Gray's 

Reef National Marine Sanctuary (MSD-05-2) 

Noise Levels and Sources in the Stellwagen Bank National Marine Sanctuary and the St. 

Lawrence River Estuary (MSD-05-1) 

Biogeographic Analysis of the Tortugas Ecological Reserve (MSD-04-1) 

A Review of the Ecological Effectiveness of Subtidal Marine Reserves in Central California 

(MSD-04-2, MSD-04-3) 

Pre-Construction Coral Survey of the M/V Wellwood Grounding Site (MSD-03-1) 

Olympic Coast National Marine Sanctuary: Proceedings of the 1998 Research Workshop, 

Seattle, Washington (MSD-01-04) 

Workshop on Marine Mammal Research & Monitoring in the National Marine Sanctuaries 

(MSD-01-03) 

A Review of Marine Zones in the Monterey Bay National Marine Sanctuary (MSD-01-2) 

Distribution and Sighting Frequency of Reef Fishes in the Florida Keys National Marine 

Sanctuary (MSD-01-1) 

Flower Garden Banks National Marine Sanctuary: A Rapid Assessment of Coral, Fish, and 

Algae Using the AGRRA Protocol (MSD-00-3) 

The Economic Contribution of Whalewatching to Regional Economies: Perspectives From 

Two National Marine Sanctuaries (MSD-00-2) 

Olympic Coast National Marine Sanctuary Area to be Avoided Education and Monitoring 

Program (MSD-00-1) 

Multi-species and Multi-interest Management: an Ecosystem Approach to Market Squid 

(Loligo opalescens) Harvest in California (MSD-99-1)