OCS Study

BOEM 2020-008

US Department of the Interior

Bureau of Ocean Energy Management

Pacific OCS Region

Cross-Shelf Habitat Suitability Modeling 

for Benthic Macrofauna
OCS Study

BOEM 2020-008

US Department of the Interior

Bureau of Ocean Energy Management

Pacific OCS Region

Cross-Shelf Habitat Suitability Modeling 

for Benthic Macrofauna

February 2020 

Authors: 

Sarah K. Henkel, Lisa Gilbane, A. Jason Phillips, and David J. Gillett

Prepared under Cooperative Agreement M16AC00014

By

Oregon State University 

Hatfield Marine Science Center

Corvallis, OR 97331 
DISCLAIMER

Study collaboration and funding were provided by the US Department of the Interior, Bureau of Ocean 

Energy Management (BOEM), Environmental Studies Program, Washington, DC, under Cooperative 

Agreement Number M16AC00014. This report has been technically reviewed by BOEM, and it has been 

approved for publication. The views and conclusions contained in this document are those of the authors 

and should not be interpreted as representing the opinions or policies of the US Government, nor does 

mention of trade names or commercial products constitute endorsement or recommendation for use.

REPORT AVAILABILITY

To download a PDF file of this report, go to the US Department of the Interior, Bureau of Ocean Energy 

Management Data and Information Systems webpage (https://www.boem.gov/Environmental-StudiesEnvData/), click on the link for the Environmental Studies Program Information System (ESPIS), and 

search on 2020-008.

CITATION

Henkel SK, Gilbane L, Phillips AJ, Gillett DJ. 2020. Cross-shelf habitat suitability modeling for benthic 

macrofauna. Camarillo (CA): US Department of the Interior, Bureau of Ocean Energy Management.

OCS Study BOEM 2020-008. 71 p.

ABOUT THE COVER

The September 2016 science team. L to R: Brittney Bair Swan, Abi Losli, Maritza Mendoza, Lenaig 

Hemery, Sarah Henkel.

ACKNOWLEDGMENTS

We appreciate the efforts of the captains and crews of the R/V Pacific Storm, R/V Ocean Star, and Bell 

M. Shimada. We thank John S. Oliver and Kamille Hammerstrom of the Benthic Lab at Moss Landing 

Marine Laboratories for sharing central California macrofauna data, Walt Nelson of the EPA for sharing 

the WEMAP data and obtaining Region 9 EPA data for northern California, and Bridgette Lorhmann and 

Sydney Harris of the EPA for providing the ODMDS data. Southern California data were provided by the 

Southern California Bight Regional Monitoring Program.
i 

Contents 

List of Figures................................................................................................................................................ ii

List of Tables................................................................................................................................................. ii

List of Abbreviations and Acronyms..............................................................................................................iii

1 Executive Summary................................................................................................................................ 1

2 Introduction to the Study......................................................................................................................... 2

3 Cross-shelf Habitat Classification........................................................................................................... 3

3.1 Purpose ......................................................................................................................................... 3

3.2 Methods......................................................................................................................................... 4

3.2.1 Study Area ......................................................................................................................... 4

3.2.2 Sample Collection and Processing.................................................................................... 4

3.2.3 Statistical Analysis ............................................................................................................. 5

3.3 Results.........................................................................................................................................10

3.4 Discussion ...................................................................................................................................20

3.4.1 Relationship to Offshore Development............................................................................23

3.4.2 Conclusions .....................................................................................................................24

4 Data Gathering and Species Selection for Region-wide Habitat Suitability Modeling .........................25

4.1 Purpose .......................................................................................................................................25

4.2 Study Area...................................................................................................................................25

4.3 Approach .....................................................................................................................................25

4.4 Data Collection and Processing ..................................................................................................27

4.5 Statistical Analyses......................................................................................................................29

4.6 Results.........................................................................................................................................29

5 Literature Cited .....................................................................................................................................33

Appendix: Additional Tables and Figures in Support of Chapter 3.............................................................38
ii

List of Figures

Figure 1. Macrofaunal grab samples (n = 404) gathered for habitat classification................................... 6

Figure 2. Three-dimensional nMDS plot of all 404 stations...................................................................... 7

Figure 3. Dendrogram developed using the 393 stations (gravel-containing stations removed prior) 

showing the multi-station clusters (groups indicated with letters in parentheses) and the 

singletons. .................................................................................................................................. 8

Figure 4. Response to depth for abundance (a), areal (sample) richness (b), H' diversity (c), and 

numerical richness (d). ............................................................................................................11

Figure 5. MDS of final 378 samples with vectors for environmental variables correlating at least 

0.4 with MDS1 (listed in Table 3). ...........................................................................................12

Figure 6. Final LINKTREE of 378 stations using depth, median grain size, and latitude to 

determine splits. ......................................................................................................................14

Figure 7. Station locations for the 14 datasets included for modeling. ...................................................26

Figure A-1. Correlations among all the environmental variables collected in the study. ...........................70

Figure A-2. Total Organic Carbon (TOC) versus depth for the 256 stations (of 378 analyzed) for 

which we had TOC data. .........................................................................................................71

List of Tables

Table 1. The 16 environmental variables for which we had data for most (249) stations. ...................... 9

Table 2. RELATE values for resemblance matrices generated from different suites of taxa. ...............11

Table 3. Correlates with the 3-D MDS axes. .........................................................................................12

Table 4. Number of stations, average physical properties, sample richness, and organism 

abundance per grab for the stations that make up each of the 12 regional-habitat 

groups as well as the averages for the outliers in each depth division. ..................................15

Table 5. The top 12 characteristic species of the 12 regional habitat groups.......................................16

Table 6. Datasets collated by Henkel lab and included in Microsoft Access Database........................28

Table 7. The 43 taxa recommended for future habitat suitability modeling. ..........................................31

Table A-1. Station data for the 404 samples considered in Chapter 3. ....................................................38

Table A-2. Abundance of all Annelids collected at the 404 stations considered in Chapter 3..................48

Table A-3. Abundance of all Crustaceans collected at the 404 stations considered in Chapter 3. ..........56

Table A-4. Abundance of all Molluscs collected at the 404 stations considered in Chapter 3. ................61

Table A-5. Abundance of all other organisms not listed above collected at the 404 stations 

considered in Chapter 3. .........................................................................................................65

Table A-6. All combinations of variables considered for the LINKTREE analysis. ...................................68
iii

List of Abbreviations and Acronyms

BOEM Bureau of Ocean Energy Management

cm centimeter

CMECS Coastal and Marine Ecological Classification Standard

CTD Conductivity, Temperature, Depth

DO Dissolved Oxygen

DOI US Department of the Interior

EFH Essential Fish Habitat

EIS Environmental Impact Statement

EMAP Environmental Monitoring and Assessment Program

EPA US Environmental Protection Agency

EUNIS European Nature Information System

FRAM Fisheries Resource Analysis and Monitoring

GIS Geographic Information System

GPS Global Positioning System

GS Grain Size

μm micrometer

m meter

mm millimeter

MDS multidimensional scaling

MLML Moss Landing Marine Laboratories

MRE Marine Renewable Energy

NCCOS National Centers for Coastal Ocean Science

NETS North Energy Test Site

NMFS National Marine Fisheries Service

NNMREC Northwest National Marine Renewable Energy Center

NOAA National Oceanic and Atmospheric Administration

NSF National Science Foundation

OCNMS Olympic Coast National Marine Sanctuary

OCS Outer Continental Shelf

ODFW Oregon Department of Fish and Wildlife

ODMDS Ocean Dredged Material Disposal Site

ONR Office of Naval Research

OSU Oregon State University

PMEC Pacific Marine Energy Center

PNW Pacific Northwest

SETS South Energy Test Site

ROV Remotely Operated Vehicle

SCCWRP Southern California Coastal Water Resources Project

SETS South Energy Test Site

SGH Surficial Geologic Habitat

TN Total Nitrogen

TOC Total Organic Carbon

USGS United States Geological Survey
1

1 Executive Summary

There is increasing interest in offshore wind development, which is likely to be sited in deeper waters

(i.e., the continental slope) on the US west coast as compared to wind installations in northern Europe 

and the US east coast or to proposed wave energy deployments. While many environmental interactions 

of marine renewable energy (MRE) deployments have been considered, a certain effect will be localized 

changes to the seabed, potentially affecting benthic invertebrates living in or on the seafloor. Previous 

work sponsored by the Bureau of Ocean Energy Management (BOEM) and conducted by Oregon State 

University (OSU) characterized seafloor habitats for benthic invertebrates on the inner to mid shelf (30 to 

130 meters [m] deep). Deeper waters are less studied, and habitat suitability models are not developed for 

macrofauna (e.g., infauna) of the outer continental shelf and slope. Thus, for this study we sampled

deeper waters (60 to 525 m) to expand the domain of the habitat characterization and of the individual 

species models. Improved models have the potential to inform regional spatial planning processes for 

future consideration of MRE projects and the necessary consultations associated with leasing (e.g., on 

Essential Fish Habitat [EFH]). Improved habitat suitability models could also improve site assessments 

needed for National Environmental Policy Act analysis and may reduce site survey requirements for 

lease holders.

The two primary outputs for this study are (1) cross-shelf habitat characterizations where benthic habitats

are classified based on biological species groupings or assemblage distributions and (2) habitat suitability 

models for a subset of the macrofauna. The cross-shelf habitat characterization was conducted for the 

same latitudinal range as the prior BOEM study and includes data from both projects as well as other 

OSU collected samples for a depth range of 20 to 525 m deep. Outputs focused on linking spatial 

variability in macrofauna assemblages with measured environmental parameters. The habitat suitability 

models were developed using the entire US west coast for the latitudinal range by including datasets from 

additional sources. To determine which macrofauna would be selected for habitat suitability modeling 

coast-wide, we determined which species were characteristic of the habitats classified and choose those 

with ranges throughout the study region. Additionally, we included species that comprise a wide range of 

tolerance/sensitivity to pollution. Finally, all seven of the species modeled previously were included in 

the final list of 44 species for which we conducted habitat suitability modeling. This study was a joint 

effort with the National Centers for Coastal Ocean Science (NCCOS). NCCOS took environmental and 

biological data from this study and built habitat suitability models. NCCOS will issue a separate BOEM 

report under Interagency Agreement Number M16PG00014.

Significant differences in macrofauna species assemblages were found across depth and in different 

sediment types. Depth was the primary structuring variable (i.e., macrofauna species assemblages first 

separated by depth) with sediment parameters (percent fines, grain size, and total organic carbon) 

secondary and nearly as statistically important. The depth break of ~90 m was the first major 

environmental variable that correlated with major differences among assemblages among our stations, 

which is not a classically described depth break for shelf fauna. Beyond the 90-m break point, we 

detected secondary changes in assemblage composition at 43 m and 200 m. Within each of these depth 

zones on the shelf, we detected significant differences in assemblages related to different sediment grain 

sizes. Below the 200-m break – a depth traditionally delineating the continental shelf and slope –

subsequent depth breaks were detected at 221 m and 445 m. Knowing how macrofaunal communities 

respond to changes in grain size and depth can inform future site surveys and led us to develop tools for 

mapping macrofauna based primarily on these physical factors.

This study provides BOEM with information on seafloor habitats and invertebrate communities in 

consideration of Outer Continental Shelf (OCS) renewable energy development. Overall, the information 

derived from this study has greatly contributed to the greater body of knowledge regarding seafloor 

habitats and invertebrate communities in the Pacific Northwest (PNW).
2

2 Introduction to the Study

The seafloor is an important resource as habitat for commercial fisheries and supports the trophic base for 

many ecologically important and regulated marine species. The vast majority of benthic habitats in waters 

past the photic zone consist of unconsolidated sediments where macrofaunal (infaunal) and epifaunal 

invertebrate species play an important role in serving as trophic links between water column and benthic 

primary productivity and higher trophic levels (Snelgrove et al. 1998; Kędra et al. 2015). A benthic 

habitat type is defined as, “a particular environment which can be distinguished by its abiotic 

characteristics and associated biological assemblage, operating at particular, but dynamic spatial and 

temporal scales in a recognizable geographic area (ICES 2006).” These invertebrates are a key part of 

defining unconsolidated sediment habitats because species represent long-term environmental conditions 

(Elliot 1994), both responding to and affecting local sedimentary processes (Gray 1974, Rhoads 1974, 

Aller and Aller 1998), benthic boundary layer flow (Friedrichs and Graf 2009), and biochemical cycling 

(Josefson and Rasmussen 2000; Laverock et al. 2014). Macrofauna-based assessment indices have been 

developed to quantify these responses and are a primary biological tool utilized worldwide to evaluate the 

overall condition or health of benthic habitats in a variety of regulatory or disturbance assessment 

contexts (e.g., waste water discharge, dredge disposal) around the world (Gillett et al. 2015; Diaz et al. 

2004).

Despite the wide-use of assessment indices in water-quality based coastal regulation, the use of 

macrofauna to classify benthic habitats as a means to inform planning applications is not always possible 

nor done consistently (Reiss et al. 2015). In US Federal waters on the continental shelf, marine planning 

primarily addresses commercial fishing, conservation, and energy development. Laws and regulations 

specifically state the importance of benthic habitats. For example, the Magnuson-Stevens Fishery

Conservation and Management Act regulates commercial ground fisheries and recognizes benthic habitat 

integration for fish species food and shelter through Habitats of Particular Concern and Essential Fish 

Habitat designations. Conservation of marine resources in the National Marine Sanctuaries on the US 

west coast covers 13,000 square miles of the seafloor. Macrofauna-focused analysis could assist with 

conservation (Bremner 2008) and inform spatial planning for groundfisheries (Thrush and Dayton 2002) 

and fish farming (Tomassetti and Porrello 2005).The Outer Continental Shelf Lands Act states the need to 

characterize and monitor benthic habitats in areas potentially impacted by energy development. Deep-sea 

coral distributions currently are utilized to delineate habitats of concern or warrant protection as these 

organisms are themselves fragile and characterize emergent bottom structure of limited extent and thus 

are consequently of increased concern. However, the much more extensive soft sediments are not 

themselves uniform, and also harbor areas which should be of concern. Prior to oil and gas leasing 

offshore southern and central California, extensive macrofauna data was collected (Hyland et al. 1991; 

Lissner 1989) but not utilized for planning purposes. In contrast, research has been conducted in the 

Alaskan arctic to link macrofauna data with higher trophic species (Kędra et al. 2015) and has informed 

energy policy in the Beaufort and Chukchi Seas.

Macrofauna assemblages represent the biological community most likely to be impacted by energy 

development. Further, the area of impact to benthic organisms could be larger than the direct footprint of 

development. Since sediment grain size often determines which animals can live in the sediment, changes 

to sediment movement due to ocean energy extraction or alterations of flow around large device arrays 

may affect the distribution of invertebrate species that are dependent on grain size, near-bottom 

sedimentation and particle loads (Etnoyer and Morgan 2003). In this study we conducted macrofauna 

community analysis and correlated it with environmental covariates as an approach for applied decisionmaking at the regional scale needed for planning applications of the US west coast seafloor.

A prior regional analysis (referenced as Benthic Habitat Characterization; BOEM-BHC, Henkel et al. 

2014) was spatially comprehensive within the mid- to inner-continental shelf (30 to 130 m) that wave 
3

energy projects have targeted – the primary focus of offshore renewables at the time. In 2018, BOEM 

issued a Call for Information and Nominations for areas with water depths from 300 to 1,200 m (Fed Reg 

2018). These deeper waters are less studied, and habitat suitability models have not been developed for 

macrofauna of the outer shelf and slope. Thus, we aimed to sample these deeper waters in order to expand 

the domain of the habitat characterization and habitat suitability models developed in the 2014 BHC 

project. Additionally, since the time of the BHC project, there has been greater interest in siting 

renewable energy projects offshore California. Thus, we aimed to expand the latitudinal range of the 

models as well by utilizing previously gathered data from various other survey programs in California as 

well as in Oregon. Evaluating and improving the spatial extent of habitat suitability models is necessary 

before they can be useful tools for siting and permitting.

The following chapters step through the study components. Chapter 3 describes box core surveys and 

subsequent analyses to describe macrofaunal invertebrate species assemblages and the classification of 

benthic habitat in the northern portion of the planning region. Chapter 4 describes the processes used for 

selecting species modeled for habitat suitability throughout the entire planning region from the CaliforniaMexico border to Vancouver, Washington.

3 Cross-shelf Habitat Classification

3.1 Purpose

New technologies are in development to produce renewable energy (wave, wind, etc.) to reduce 

dependence on fossil fuels. The energetic continental shelf of the PNW is particularly appealing for 

marine renewable energy development and areas of the seafloor have the potential to be leased to 

developers for this purpose. An assessment of potential impacts on marine environments is needed, but 

the lack of baseline data on the characteristics, distribution, abundance and condition of seabed habitats 

limits our ability to predict how they might change. However, even before seabed habitat distribution, 

abundance and condition can be assessed, a better approach for classification of seabed habitats is 

necessary. In the rationale for Session G: Habitat Modelling and Mapping for better assessment and

monitoring of our seas at the ICES Annual Science Conference in 2011, conveners stated, “empirical 

evidence shows this new categorisation [EUNIS] results in a poor match between modelled and observed 

biotopes; primarily because the boundaries between classes have not been defined because of any known 

effect on benthic community distribution”. This knowledge gap also affects other aspects of marine 

spatial planning, including evaluation of potential impacts on biological resources such as fisheries and 

other higher trophic levels that may respond to the distribution of benthic communities rather than 

physical features.

Due to their living position and life history, macrobenthic fauna are both indicators of sediment 

characteristics and sensitive to changes to sediment conditions; thus, they are the biological community 

most likely to be impacted by seafloor development. Not only do they serve as a food source for higher 

trophic levels, they also are a link between toxicants in the sediments and bioaccumulation in tissues of 

fishes, marine mammals, birds, and humans. Thus, benthic macrofauna have long been used as indicators 

of both habitat and environmental status and trends. Planning for both offshore renewable energy and 

fisheries management is ongoing across the shelf and slope-shelf transition zone in the PNW, driving the 

need for better understanding of the distributions of habitats as it relates to these critical, low trophic level 

species.

In southern California, where there are extensive long-term macrobenthic fauna-based regional 

monitoring programs in place across the continental shelf and slope (Schiff et al. 2016), depth has been

considered to be the primary variable structuring macrofaunal invertebrate species distributions with other 
4

factors such as dissolved oxygen, grain size, and total organic carbon being secondary (SAIC 1986, 

Lissner 1989, Hyland et al. 1991, Bergen et al. 2001, Gillett et al. 2017). Henkel and Politano (2017) 

focused on the mid-continental shelf (50 to 110 m) macrofauna of the PNW and defined important breaks 

in sediment characteristics that resulted in different macrofaunal assemblages within that depth range in 

the region. Across a broader latitudinal (US west coast) range and slighter greater depth expanse (30 – 130 

m), Henkel and Nelson (2018) similarly found within-region differences based on low percentages of fine 

sediment along with latitudinal breaks in species assemblages, potentially related to temperature and/or 

upwelling. However, because of the relatively limited depth range in both studies, we were not able to 

determine if/where significant depth breaks in macrofaunal assemblages occur across the entirety of the 

shelf and slope, if those breaks are consistent with those observed in Southern California, or if the 

sediment classifications defined in the previous studies apply across the broader geographic scope of the 

outer shelf and slope of the PNW.

The purpose of this study was to understand how macrofauna are distributed in relation to physical factors

across the shelf and the upper slope (~20 to 525 m) from Mendocino, California, to Grays Harbor, 

Washington. The objective was to classify sedimentary benthic habitats as defined by the macrofaunal

organisms. We specifically sought to determine (1) where significant depth-related breaks in species 

assemblages occurred; (2) what sediment differences correlate with different species assemblages; and 

(3) if regional differences were detectable in this sampled range. The results of this effort will more 

accurately characterize a large seafloor expanse and assess the impacts of marine renewable energy 

development.

3.2 Methods

3.2.1 Study Area

The study area ranged from Fort Bragg, California (39.5 °N) to Grays Harbor, Washington (47.0 °N) on 

the west coast seafloor of the continental United States. From 2010 to 2012 a total of 242 samples were 

collected from nine distinct sites in this range; within each site, sample stations were randomly 

determined using Generalized Random Tessellation Stratified methods (GTRS; Stevens and Olsen 2004) 

as described in Henkel and Politano (2017). In early summers 2014 and 2015, we collected 77 samples 

from 60 to 500 m deep along regularly spaced (0.5 degrees latitude) offshore transects from Brookings to 

the Columbia River, Oregon. An additional site near Newport, Oregon, was sampled in June of 2015, 

within which 17 stations were arranged on a regular grid. In summer 2016, we sampled 68 additional 

stations from Coos Bay to Tillamook, Oregon, from 64 to 525 m deep. Thus, a total of 404 samples 

ranging from 18 to 525 m deep were available for analysis (Figure 1; Table of all station locations in the 

Appendix).

3.2.2 Sample Collection and Processing

At each station, samples were collected with a modified Grey-O’Hara 0.1 m2 box core. One box core 

sample was taken at each station. Only samples with a penetration depth of at least 5 centimeters (cm) and 

no slumping or other evidence of disturbance (i.e., by washing) were accepted for processing. 

Approximately 80 milliliters (mL) of sediment were collected from the undisturbed surface layer for grain 

size analysis. Any organisms noticed in the sediment subsample were removed and placed in the 

organism sample at the time of collection (occasionally a specimen was detected when conducting the 

sediment analysis). The remainder of the collected core was sieved onboard through a 1.0 millimeter 

(mm) mesh screen, and all collected organisms (both infauna living in the sediment and small epifauna 

which may have been on the surface – hereafter collectively called macrofauna) were preserved in a 

mixture of 10% buffered formalin and seawater. At most stations (primarily excluding the 2014 and 2015 

offshore sampling) vertical water-column profiles of conductivity, temperature, dissolved oxygen, pH, 
5

and fluorescence were obtained with a Sea-Bird Electronics CTD (conductivity, temperature, depth) unit 

equipped with additional sensors. Depth was recorded from the vessels’ echosounder at the time the box 

corer hit the bottom.

Upon return to the laboratory, organisms were transferred to 70% ethanol then sorted into major 

taxonomic groups by OSU staff. Crustaceans, polychaetes, other worm-like creatures, and a portion of the 

molluscs were sent to contracted taxonomic experts. OSU laboratory staff identified other molluscs, 

echinoderms, and any remaining taxa.

Sediment from the top of the core was analyzed using a Beckman Coulter Laser Diffraction Particle Size 

Analyzer (LD-PSA) to determine mean and median grain size and percent fines (silt/clay; portion less 

than 62.5 micrometer [μm]; Wentworth 1922). In most cases, sand (62.5 μm to 2 mm) was the balance of 

the sample. Where grain sizes larger than 2 mm (maximum size for the LD-PSA) were encountered, these 

samples were fractioned and the percent gravel (that fraction greater than 2 mm) was determined by 

weight. The balance of the fraction was then analyzed by the LD-PSA to determine % sand and silt/clay 

and grain size (the mean/median grain sizes are thus not representative of the full composition of the 

sediment). Approximately one quarter (n = 110) of the sediment samples were analyzed for percent total 

organic carbon (TOC) and total nitrogen (TN) using an NA1500 Elemental Analyzer operated by staff in 

the College of Earth, Ocean, and Atmospheric Sciences at OSU.

3.2.3 Statistical Analysis

A variety of multivariate analyses were undertaken using PRIMER 6th Edition (Clarke and Gorley 2006). 

Abundance data were 4th root transformed to reduce the influence of highly abundant species without 

giving as much weight to rare species as would the more severe log(x+1) (Clarke and Warwick 2001). A 

Bray-Curtis similarity matrix was created for the 404 samples and a multidimensional scaling (MDS) plot 

was created. As Henkel and Politano (2017) determined that stations containing gravel and those with less 

than 1% fines hosted significantly different assemblages, and Henkel and Nelson (2018) detected a 

significant difference in assemblages at a break of 5% fines, we first coded the MDS with sediment 

classes representing these and other breaks in percent fines (Figure 2). As expected, the gravel-containing 

stations were rather separate from the rest of the stations. These stations represent a unique habitat type 

with its own assemblage of organisms (characterized mostly by polychaetes; Henkel and Politano 2017); 

however their inclusion in this analysis would make other differences within gravel-free sediment 

difficult to resolve; thus, we removed all 11 stations that contained gravel. Additionally, we removed 

from the dataset any taxon that had only a single observation. A new resemblance matrix was created 

using the remaining 393 samples. We used the similarity matrix to create a dendrogram by agglomerative 

cluster analysis with group-average linking and the Similarity Profiles (SIMPROF) procedure to identify 

significant clusters based only similarities in macrofaunal abundance. Using a SIMPROF significance 

threshold of 1%, the dendrogram included 76 multi-station clusters and 15 individual samples as outliers 

(Figure 3). These outliers were removed from the remainder of the analysis for a final dataset of 378 

samples.
6

Figure 1. Macrofaunal grab samples (n = 404) gathered for habitat classification.

The base map is the primary lithology as described in the BOEM-BHC project (Goldfinger et al. 2014).
7

Figure 2. Three-dimensional nMDS plot of all 404 stations.

Gravel stations are grey boxes on the left side of the plot.
8

Figure 3. Dendrogram developed using the 393 stations (gravel-containing stations removed prior) showing the multi-station clusters

(groups indicated with letters in parentheses) and the singletons.

Significance evaluated at the 1% level found 76 distinct groups and 15 singletons, which were removed for final analysis.
9

A final resemblance matrix of the 378 stations was created for the entire suite of taxa, and individual 

matrices were created for polychaetes, molluscs, crustaceans, and echinoderms. These resemblance 

matrices were then compared between all pairs of sub-groups, and sub-groups were compared to the 

matrix based on all taxa using the RELATE procedure in PRIMER to determine if patterns were similar 

for the different subgroups and which subgroup was most similar to the matrix generated with all taxa.

The final set of 378 stations were plotted on 2-D and 3-D MDS planes and potential physical correlates to 

MDS axes were assessed. Here we included all station and sample data [Latitude, Depth, sediment 

fraction less than 62.5 μm (Fines%), Mean Grain Size, Median Grain Size, total organic carbon (TOC), 

total nitrogen (TN), and bottom values from CTD casts (Temperature, DO, Salinity, Fluorescence, 

Turbidity, Beam Transmission, and pH)]. We also included upwelling indices from NOAA’s Pacific 

Fisheries Environmental Laboratory 

(https://www.pfeg.noaa.gov/products/PFELData/upwell/monthly/upindex.mon) for the month of 

sampling (UpwellingMon), the previous month (UpwellingPrev), and the average for the year 

(UpwellingYear) for the latitudinal zone of each of the stations. Those parameters with greater than 

R = 0.4 to any MDS axis were plotted on the included 2-D plot.

To determine which measured environmental factors “best” described the distribution of macrofaunal 

invertebrates across the study region, the BIO-ENV (BEST) procedure was used to find the subset of 

environmental variables (summarized as pairwise normalized Euclidean distance) with the strongest 

Spearman rank correlations to the macrofauna observed among the samples (summarized as pairwise 

Bray Curtis similarities). The BEST procedure requires a complete inventory of biological and 

environmental data at all stations in the two similarity matrices for analysis; thus, only Latitude, 

Longitude, Depth, Fines%, Mean Grain Size, Median Grain Size, UpwellingMon, UpwellingPrev, and 

UpwellingYear could be considered in the analysis of all stations. We additionally conducted a BEST 

analysis using only the stations (n = 249) for which we had the majority of the environmental data (16 

variables; Table 1). In both cases environmental data were normalized prior to running the procedure.

Table 1. The 16 environmental variables for which we had data for most (249) stations.

Variables in italics are those for which we had data for all (378 stations).

Station Data Sediment Data CTD Data Indices

Latitude Fines% Temperature UpwellingMon

Longitude MeanGS Dissolved Oxygen UpwellingPrev

Depth MedianGS Salinity UpwellingYear

TOC Fluorescence

TN pH

The variables determined to be highly correlating with the species assemblage patterns were then used as 

the starting point to classify the stations in the LINKTREE procedure in PRIMER 6. The LINKTREE 

procedure is a form of constrained cluster analysis involving a divisive partition of the biotic community 

samples into ever smaller groups, but in which each division has an 'explanation' in terms of a threshold 

on one of the environmental variables (Clarke et al. 2008). We constrained the minimum group size to be 

two stations (so as not to have singletons) and used the SIMPROF procedure (using the Bray-Curtis 

similarity index) to finalize the tree when there were no significant differences (p < 0.01) among the 

remaining stations. Within a resulting LINKTREE, each branch of the tree corresponds to a group of 

samples with similar macrofaunal composition (based on taxon and abundance); each split maximizes the 

Analysis of Similarity (ANOSIM) R statistic between macrofaunal groups and reports the threshold value 

of the environmental variable associated with that division (Clarke et al. 2008). If a threshold in more 
10

than one environmental variable corresponds to a break between groups of stations in the LINKTREE, 

threshold values for both variables are reported. We wanted to explore which combination of physical 

variables yielded the most parsimonious LINKTREE (i.e., the fewest number of splits). Thus, we created 

LINKTREEs using each of the physical parameters individually, in pairs, in trios, in groups of four, and 

with all the potential predictors.

Additionally, abundance, areal taxa richness, and Shannon-Weiner diversity were calculated from the 378 

stations (samples) and plotted against depth to summarize how macrofaunal abundance and diversity 

changed across the shelf and the upper slope and the significance of that relationship was tested using 

linear regression. Abundance equals the number of all animals per station, areal (sample) richness is the 

number of taxa per box core (0.1 m2 area of seafloor), and numerical richness is the number of taxa per 

number of individuals at a station. Shannon’s diversity index (H' diversity) weights the geometric mean of 

the proportional abundances of the taxon groups.

3.3 Results

The final dataset for analysis of 378 stations ranged from 18.2 m deep to 525 m deep and from 39.5° to 

47° N. Actual penetration depth of the box corer averaged 21.3 cm with a median of 23.75 cm for an 

average total core volume of over 2000 cm2

.

The highest correlations among environmental parameters were between percent fine sediment and mean 

and median grain size (-0.76/-0.77), as expected as they are different measures of the same component. 

The second highest correlations were between TOC/TN and percent fine sediment (0.84/0.81) followed 

by TOC/TN correlations with depth (0.78/0.79). Latitude correlated most strongly with upwelling (as we 

based the upwelling indices on the latitude of the stations) and secondarily with percent fines (-0.60). 

Correlates among all environmental variables are given in Figure A-1 in the Appendix.

Organism abundances ranged from 10 to 542 individuals per 0.1 m2 grab, and sample richness ranged 

from 5 to 64 taxa per grab. In total, 757 taxa were identified and included for analysis, most to the species 

level. Seven of the top ten most abundant species in the entire dataset were bivalves with two polychaetes 

and the pea crab, Pinnixa occidentalis complex, making up the other three (see Tables A-2 through A-5 in 

the Appendix). Overall, both organism abundance and sample richness (per 0.1 m2

) declined significantly 

(p < 0.001) with depth; however, no significant response to depth was detected for Shannon’s diversity 

index (p = 0.753) while numerical species richness (number of taxa per number of individuals per grab) 

significantly increased (p < 0.001) with depth (Figure 4). The resemblance matrix generated with the full 

suite of taxa was most similar to molluscs only with a Rho of 0.828 and secondarily similar to the 

polychaetes only with a Rho of 0.751. The resemblance matrices of the different groups compared 

amongst each other were not highly related (Table 2).
11

Figure 4. Response to depth for abundance (a), areal (sample) richness (b), H' diversity (c), and 

numerical richness (d).

Table 2. RELATE values for resemblance matrices generated from different suites of taxa.

Taxon Group Taxon Group RELATE Value

All taxa Molluscs 0.828

All taxa Polychaetes 0.751

All taxa Crustaceans 0.461

All taxa Echinoderms 0.377

Polychaetes Molluscs 0.392

Polychaetes Crustaceans 0.269

Polychaetes Echinoderms 0.230

Molluscs Crustaceans 0.304

Molluscs Echinoderms 0.200

Crustaceans Echinoderms 0.127
12

The MDS indicated a separation of assemblages with less than 2% fines, with 2-5% fines intermediate 

between those low silt containing stations and the remainder. Stations with >5% fines did not appear to 

further separate based on Folk (1974) defined breaks in silt content (Figure 5). Depth was the highest 

correlate with MDS1 (r = -0.725 for 2-D) with sediment grain size parameters the second highest 

correlate with MDS1. Stress was quite high for the 2-D MDS, and there were no correlates higher than 

0.5 with the second MDS axis. Thus, the correlates to the 3-D MDS were determined and reported

(Table 3). Depth and sediment grain size parameters remained the highest correlates of MDS1, followed 

by CTD parameters and the upwelling index for the previous month. Depth (again) and dissolved oxygen 

were the highest correlates of MDS2. All correlations of environmental variables with MDS3 were low.

Figure 5. MDS of final 378 samples with vectors for environmental variables correlating at least 0.4 

with MDS1 (listed in Table 3).

Stations are plotted according to the Folk (1974) percent fines classification with the addition of <1% and 1-2% fines 

based on the findings of Henkel and Politano (2017).

Table 3. Correlates with the 3-D MDS axes.

Variables were included in the table if they correlated >0.4 with any axis on the 3-D MDS plot. Bolded values on 

MDS1 are included on Figure 5.

Depth Fines%

Mean 

GS

Median 

GS TOC TN Temp DO Salinity pH

Upwell 

Prev

MDS1 0.722 0.702 -0.684 -0.695 0.446 0.424 -0.468 -0.412 -0.414 -0.488 0.472

MDS2 -0.502 -0.125 -0.221 -0.217 -0.111 -0.143 0.238 0.362 0.124 0.117 -0.028

MDS3 0.162 -0.268 0.055 0.035 -0.196 -0.177 -0.321 -0.176 -0.268 -0.283 -0.201
13

The BEST match between the biological data and the environmental data for which we had complete 

coverage used just two variables: depth and median grain size, with a rather high correlation (r = 0.643); 

however, we could not take any water column parameters or sediment TOC/TN into account in the full 

BEST analysis because we did not have data for all stations. Using just the stations (n = 249) for which 

we had the majority of environmental data (Table 1) the highest correlation used depth, mean grain size, 

median grain size, TOC, and fluorescence with r = 0.655 (using just depth and median grain size with the 

subset of stations resulted in r = 0.637). Despite being included in the BEST results, fluorescence’s 

highest correlation with any MDS axis was just -0.283 with MDS1.

The initial LINKTREE run using only depth and median grain size resulted in 101 splits. Adding latitude 

resulted in a LINKTREE with 71 splits: the lowest number of splits with the fewest variables (see Table 

A-6 in the Appendix for all options) and only two more than using all the physical parameters, for a total 

of 72 distinct groupings of stations (Figure 6). Among the 71 splits, four minor splits (F, K, Q, AH) could 

be described by more than one of the three environmental variables. ANOSIM of these 72 groups had a 

global R of 0.892. We grouped the 72 termini of the LINKTREE into 12 major habitats based on a cut-off 

of 25% separation, plus one further major sediment split at B=18.6% (separating groups IV and V). We 

did not include splits with higher B% that resulted in < 5 stations on a terminus as we deemed those more 

outliers rather than separate habitats. Analyzing these 12 habitat types using ANOSIM resulted in a global 

R of 0.792.

The average values of the physical parameters that correlated with the MDS axes are given in Table 4 for 

each habitat group. Also shown are the average number of species and the average number of organisms

per 0.1 m2 box core grab. The individual average abundance per 0.1 m2 box core and the cumulative 

percentage contribution to within group similarity of the top 12 “characteristic” species (those species 

whose abundances are highly contributing to the average similarity within a group as determined by 

SIMPER) of the 12 regional habitat groups determined by the LINKTREE analysis are shown in Table 5.
14

Figure 6. Final LINKTREE of 378 stations using depth, median grain size, and latitude to determine splits.

Breaks that were used to determine the 12 regional-habitat groups are detailed in blue.

Inner and Mid Shelf

Groups I to V

Outer Shelf and Upper Slope Groups VI 

to XII
15

Table 4. Number of stations, average physical properties, sample richness, and organism abundance per grab for the stations that make 

up each of the 12 regional-habitat groups as well as the averages for the outliers in each depth division.

Supervised Group N

Depth 

(m)

Fines 

%

Mean 

GS

Median 

GS TOC TN

Temp 

(°C)

DO 

(ml\L) Salinity pH

Avg. 

Rich

Avg. 

Abund.

I: Mid-Shelf

 Muddy NorCal 13 65.2 68.4 53.2 48.7 0.72% 0.07% 9.01 3.10 33.72 7.81 41.7 167.2

II: Inner Shelf

 (Clean Sand) 41 33.8 0.5 256.6 236.0 0.07% 0.01% 8.01 2.52 33.66 7.73 21.2 68.6

III: Mid Shelf

 <223 um 48 69.6 7.8 185.3 175.7 0.31% 0.03% 7.58 1.90 33.79 7.69 37.3 195.1

IV: Mid Shelf

 <301 um 46 63.7 0.4 286.8 251.4 0.08% 0.01% 7.71 1.69 33.76 7.67 26.3 179.6

V: Mid Shelf

 >315 um 33 63.9 0.4 459.6 430.1 0.07% 0.01% 7.79 1.68 33.78 7.73 20.8 94.1

VI: Outer Shelf

 <104 um North 15 161.5 50.0 72.4 68.2 1.38% 0.14% 7.28 1.78 33.82 7.71 30.9 103.9

VII: Outer Shelf

 <104 um South 59 114.9 63.2 55.4 48.3 0.91% 0.10% 8.31 2.02 33.68 7.72 27.7 111.0

VIII: Outer Shelf

 111-167 um 8 122.3 9.3 152.5 145.9 0.59% 0.06% 7.49 1.83 33.88 7.72 47.4 184.4

IX: Outer Shelf

 >170 um 17 100.2 3.5 230.5 205.8 0.26% 0.03% 7.49 1.81 33.82 7.72 43.4 147.5

X: Slope Break 13 207.4 38.7 144.9 118.5 1.24% 0.14% 7.30 2.35 33.96 7.88 29.2 96.5

XI: Upper Slope 48 336.1 47.5 90.1 76.5 1.47% 0.16% 6.42 1.45 34.02 7.78 23.7 75.8

XII: Slope 13 484.2 80.9 39.0 28.4 1.62% 0.17% 5.28 0.69 34.13 7.66 13.6 25.7

Inner Shelf Outlier 4 19.0 0.55 197.6 193.3 n.d. n.d. 7.84 2.13 33.67 7.70 21.3 106.0

Mid Shelf Outlier 7 70.2 0.8 508.6 479.4 0.12% 0.02& 7.63 1.35 33.83 7.65 17.9 60.1

Outer Shelf Outlier 4 124.5 16.6 325.3 272.2 n.d. n.d. 7.68 2.31 33.87 7.72 15.0 48.3

Slope Outlier 9 354.1 65.6 96.7 73.2 2.46% 0.29% 6.56 1.51 34.03 7.81 22.2 57.2
16

Table 5. The top 12 characteristic species of the 12 regional habitat groups.

Species and percent characteristic (Cum. %) were determined using SIMPER. Average abundance per 0.1 m2 box 

core in each group and the cumulative percentage contribution to within group similarity are listed.

I. Mid-Shelf Muddy NorCal Avg. similarity: 41.15

Av. Abund Cum.%

Axinopsida serricata 33.2 9.62

Ennucula tenuis 16.3 18.12

Ninoe gemmea 4.3 23.21

Ampelisca careyi 2.7 28.25

Magelona longicornis 1.5 32.69

Macoma spp 1.7 36.72

Sternaspis fossor 3.2 40.69

Acteocina spp 1.4 44.56

Euclymeninae spp 0.9 48.09

Turbonilla spp 0.8 51.62

Paraprionospio alata 0.8 55.12

Maldane sarsi 1.2 57.93

II. Inner Shelf Clean Sand Avg. similarity: 30.60 III. Mid Shelf <223 um Avg. similarity: 31.78

Av. Abund Cum.% Av. Abund Cum.%

Callianax pycna 1.6 9.40 Axinopsida serricata 37.2 14.51

Carinoma mutabilis 1.0 17.88 Ennucula tenuis 2.4 20.85

Scoloplos acmeceps 0.7 24.60 Pinnixa occidentalis complex 3.0 25.72

Majoxiphalus major 0.6 30.58 Acila castrensis 1.6 30.60

Dendraster excentricus 0.3 35.74 Amphiodia urtica 0.7 34.05

Chaetozone bansei 0.3 40.41 Onuphis iridescens 0.4 37.17

Nephtys caecoides 0.2 44.91 Kurtiella tumida 1.4 40.24

Rhepoxynius vigitegus 0.4 48.96 Scoletoma luti 0.5 42.91

Magelona sacculata 0.2 53.00 Ampelisca careyi 0.3 45.56

Eohaustorius sencillus 0.3 56.73 Nephtys spp 0.2 48.07

Rhepoxynius spp 0.2 60.42 Cylichna attonsa 0.2 50.56

Eohaustorius sawyeri 0.2 63.84 Rhepoxynius spp 0.2 52.86

IV. Mid Shelf <301 um Avg. similarity: 39.72 V. Mid Shelf >315 um Avg. similarity: 35.22

Av. Abund Cum.% Av. Abund Cum.%

Axinopsida serricata 32.6 15.01 Nutricola lordi 10.0 14.78

Nutricola lordi 29.0 27.73 Ophelia assimilis 3.2 26.59

Cylichna attonsa 4.2 37.10 Spiophanes norrisi 1.1 34.13

Magelona sacculata 1.2 42.57 Tellina nuculoides 1.0 41.23

Ampelisca careyi 1.0 47.88 Axinopsida serricata 1.9 48.28

Euphilomedes carcharodonta 0.9 52.26 Callianax baetica 0.6 54.40

Callianax baetica 0.8 56.53 Cylichna attonsa 0.4 58.66

Spiophanes norrisi 0.4 59.99 Aphelochaeta spp 0.5 62.77

Onuphis iridescens 0.5 63.36 Axiothella rubrocincta 0.3 66.80

Alia gausapata 0.6 66.56 Alia gausapata 0.2 69.94

Glycinde armigera 0.3 69.69 Microspio pigmentata 0.1 72.52

Carinoma mutabilis 0.2 71.88 Pinnixa occidentalis complex 0.1 74.65
17

VI. Outer Shelf <104 um South Avg. similarity: 32.38 VII. Outer Shelf <104 um North Avg. similarity: 33.42

Av. Abund Cum.% Av. Abund Cum.%

Axinopsida serricata 14.5 13.32 Axinopsida serricata 17.7 13.92

Acila castrensis 3.3 22.16 Rhabdus rectius 4.8 22.73

Macoma carlottensis 2.9 30.88 Adontorhina cyclia 1.7 29.79

Amphioplus strongyloplax 1.0 37.22 Pectinaria californiensis 1.7 36.61

Onuphis iridescens 0.7 43.30 Galathowenia oculata 2.6 42.43

Adontorhina cyclia 0.5 47.50 Euclymeninae spp 0.5 46.18

Rhabdus rectius 0.5 51.15 Macoma carlottensis 0.5 49.92

Brisaster latifrons 0.2 53.99 Onuphis iridescens 0.4 53.60

Ninoe gemmea 0.2 56.69 Heterophoxus spp 0.5 57.13

Sternaspis fossor 0.2 59.32 Amphioplus macraspis 0.2 60.16

Paraprionospio alata 0.1 61.93 Ninoe gemmea 0.2 62.98

Ennucula tenuis 0.2 64.11 Amphiodia urtica 0.1 65.55

VIII. Outer Shelf 111-167 um Avg. similarity: 33.37 IX. Outer Shelf >170 um Avg. similarity: 31.08

Av. Abund Cum.% Av. Abund Cum.%

Spiochaetopterus costarum 12.5 6.23 Pista estevanica 1.9 5.19

Paraprionospio alata 1.5 11.97 Acila castrensis 1.5 9.98

Axinopsida serricata 6.2 17.65 Amphiodia urtica 1.1 14.70

Magelona longicornis 3.3 22.77 Axinopsida serricata 2.6 19.12

Rhabdus rectius 3.1 27.53 Onuphis iridescens 1.0 23.36

Amphioplus macraspis 2.0 32.28 Cylichna attonsa 1.0 27.55

Nephtys ferruginea 1.2 36.91 Magelona berkeleyi 1.6 31.52

Paradiopatra parva 1.3 41.22 Ennucula tenuis 1.5 35.36

Euclymeninae spp 1.1 45.13 Prionospio steenstrupi 1.6 39.13

Myriochele gracilis 1.8 48.53 Glycera nana 0.4 42.26

Thyasira flexuosa 0.8 51.85 Huxleyia munita 0.8 45.18

Glycinde armigera 0.4 54.68 Paraprionospio alata 0.4 48.01
18

X. Slope Break (200 – 221 m) Avg. similarity: 24.27 XI. Upper Slope (221–445 m) Avg. similarity: 29.33

Av. Abund Cum.% Av. Abund Cum.%

Axinopsida serricata 3.8 13.04 Huxleyia munita 8.8 16.68

Adontorhina cyclia 2.4 25.59 Galathowenia oculata 1.9 25.29

Galathowenia oculata 4.4 33.70 Gadila tolmiei 0.9 33.74

Heterophoxus spp 0.3 39.18 Onuphis iridescens 0.8 41.78

Onuphis iridescens 0.5 43.80 Brisaster latifrons 0.4 48.61

Sternaspis assimilis 0.2 48.08 Ennucula tenuis 0.4 54.89

Rhabdus rectius 0.2 52.26 Chaetoderma argenteum 0.4 60.69

Maldane sarsi 0.2 56.32 Rhabdus rectius 0.2 64.65

Aricidea (acmira) simplex 0.2 59.26 Adontorhina cyclia 0.2 67.97

Amphioplus macraspis 0.1 62.18 Amphioplus macraspis 0.1 70.76

Huxleyia munita 0.1 64.22 Axinopsida serricata 0.1 73.41

Mendicula ferruginosa 0.1 66.25 Pectinaria californiensis 0.1 75.86

XII. Slope (445 – 525 m) Avg. similarity: 28.05

Av. Abund Cum.%

Huxleyia munita 1.00 17.16

Rhabdus rectius 0.50 29.31

Maldane sarsi 0.70 40.54

Chaetoderma argenteum 0.30 49.52

Glycinde armigera 0.10 56.33

Phoronidae sp 0.20 61.89

Gadila tolmiei 0.06 66.47

Brisaster latifrons 0.04 70.27

Cerebratulus spp 0.03 73.44

Praxillella gracilis 0.03 76.27

Eucranta anoculata 0.03 79.08

Artacama coniferi 0.03 81.76
19

The first major split in the LINKTREE (C) was associated with a depth break at 89 m. We classified the

shallower side of this split (left side of the tree) as mid and inner shelf habitats. Within this group, the 

next split (D) was a latitude split at 42°, which is essentially the Oregon-California (OR-CA) border. 

Stations on the left side of this split were from the site off Eureka, California, except for one: the 

southern-most station sampled in Oregon. This “Mid-Shelf Muddy NorCal” group (Group I) of stations 

ranged 52 to 77 m deep and had much higher percent fine sediment (68.4%) than the rest of the mid to 

inner shelf stations (average 2.5%). The mid-shelf, muddy northern California community was 

characterized by the ubiquitous bivalves Axinopsida serricata and Ennucula tenuis. Unique to Group I 

were the polychaetes Ninoe gemmea, Magelona longicornis, and Sternaspis fossor, which are uncommon 

on the sandier, mid-shelf stations of Oregon.

The next split (G) define a break between the inner- and mid-shelf. Stations on the shallower side of this 

split ranged 18.2 to 42.5 m deep and consisted mostly of clean sand (0 – 1.82% fines) with average 

median grain size of 235 μm; thus, we called this “Inner Shelf Clean Sand” (Group II). The top 

characteristic species of this group were the snail Olivella (Callianax) pycna, the nemertean, Carinoma 

mutabilis, the polychaete, Scoloplos acmeceps, the amphipod, Majoxiphalus major, and the sand dollar, 

Dendraster excentricus, of which only the polychaete was characteristic in any other habitats. Notably, no 

bivalves were in the top 12 characteristic species for the Inner Shelf Clean Sand habitat. A subsequent 

split in this group was detected at 44.5° latitude with a separation value of 18.9%.

On the mid-shelf (depths 42.8 – 88 m) the next split (N) was at </>223 μm. On the mid-shelf where grain 

sizes were less than 223 μm (left side of split N, Group III), stations averaged 7.8% fines and hosted 

slightly different assemblages in the north and south of the study region (44.0 °N split P, with a similarity 

of 18.6%, similar in separation value and latitude as found in Group II). As in California at these depths, 

the bivalves A. serricata and E. tenuis were highly characteristic, and these regions shared three other 

characteristic species. Additionally, the pea crab, Pinnixa occidentalis complex, and the bivalves, 

Kurtiella tumida and Acila castrensis were highly contributing (particularly in the south) as well as the 

brittle star Amphiodia urtica and Onuphis iridescens (particularly in the north). Larger than 223 μm, 

stations were further subdivided into <301/>315 μm (split Y, Group IV and V). Here, bivalves were 

abundant, particularly Nutricola lordi and the ubiquitous A. serricata, along with the gastropods Cylichna 

attonsa, Callianax baetica, and Alia gausapata. The larger grain sized group (Group V) had the bivalve 

Tellina nuculoides as the 4th most characteristic species, which was uncommon among stations in the 

<301 μm group (Group IV) and A. serricata was far less abundant in this grain size class than anywhere 

else on the mid shelf. Most differences between Groups IV and V were in the characteristic polychaetes: 

only Spiophanes norrisi was shared.

The deeper waters (right side of C) were structured similarly to the inner and mid shelf groups, having

three grain size groups and lower separation latitudinal breaks within four Outer Shelf and three slope 

groups. The four outer shelf groups (89 to 200 m and left side of AO) are defined by grain size. The 

smaller grain size stations (< 104 μm) split into a southern (Group VI) and northern (Group VII) groups 

(AQ at a latitude of 43.6°). This latitudinal break is similar to the break found in the mid-shelf within 

Group III at 44.0° and inner-shelf Group II at 44.5°. At these smaller grain size (< 104 μm) stations, 

bivalves Macoma carlottensis and Adontorhina cyclia (both regions) and Acila castrensis (in the south, 

Group VI) were more characteristic than the less abundant, E. tenuis. Amphioplus spp. brittle stars 

(different species in north and south) replaced A. urtica, and the heart urchin Brisaster latifrons was 

characteristic of the southern group (Group VI). The scaphopod, Rhabdus rectius was characteristic of 

both groups. Within the southern group (Group VI), a second latitudinal break was detected at 40.9°

latitude (isolating the California stations from southern Oregon) with a Similarity (B) value of 19.1%. In 

the Outer Shelf > 111 μm group there was a secondary split at <167/>170 μm (BC forming Groups VIII 

and IX). In the Outer Shelf 111-167 μm group (Group VIII) polychaetes were the most highly 
20

contributing species along with A. serricata, R. rectius, and A. macraspis. In the largest grain size class on 

the outer shelf (>170 μm, Group IX), a different suite of polychaetes was characteristic, along with,

Amphiodia urtica, Acila castrensis and Cylichna attonsa, all of which are characteristic of mid-shelf 

stations with similar grain size (Group III).

Split AO represents the 200 m shelf-slope break. The next split (BJ) isolated stations 445 m to 525 m with 

no subsequent distinctions based on sediment characteristics or latitude (although we only sampled 

stations that deep from 43.40°N to 43.75 °N). The Slope assemblage (Group XII) had only 28.05% within 

group similarity and was characterized primarily by the bivalve, Huxleyia munita, and the scaphopod, 

Rhabdus rectius. Polychaetes Maldane sarsi and Glycinde armigera and the “glistenworm” Chaetoderma 

argenteum, a shell-less mollusc, also were in the top five characteristic species. Split BN then isolated 

stations less than 217 m deep on the slope. These Slope Break stations (Group X) had the lowest within 

group similarity (24.27%; constituting three separate groups on the LINKTREE) and were characterized 

by both common shelf and slope taxa. A latitudinal break at 44 °N (BR) – as on the shelf – was detected 

in the remaining 56 Upper Slope stations between 224 and 445 m deep. Collectively, the Upper Slope 

stations were characterized primarily by the bivalve, Huxleyia munita, polychaetes, Galathowenia oculata

and Onuphis iridescens and scaphopods Gadila tolmiei and Rhabdus rectius (particularly in the south). 

The bivalves Ennucula tenuis and Adontorhina cyclia, the heart urchin, B. latifrons, and the glistenworm 

C. argenteum also were characteristic.

3.4 Discussion

Establishing an understanding of benthic species-habitat associations over broad spatial scales is useful to 

marine spatial planning for many purposes including the siting and evaluating offshore renewable energy 

projects. Just as isolated mapping data was not sufficient for the groundfish EFH process, knowing that 

100 worms were found in sand at one specific site does not alone provide the tools for resource managers 

to assess benthic resources or describe potential impacts from human activities on the sea floor. However, 

knowing the distributions of benthic invertebrates at a regional scale provides data applicable to 

cumulative impact assessments and context for project-specific surveys.

We sought to classify benthic sedimentary habitats across the shelf and upper slope in the PNW based on 

the macrofaunal organisms living in and on the sediment. This and a previous BOEM study (Henkel et al. 

2014) represent the first attempts to examine spatial variability in macrofaunal communities with 

consistent sampling and identification methodology from the inner shelf, across the shelf break, and down 

the slope in the eastern north Pacific.

Previous studies in the region focused only on the mid to outer shelf, and did not find major distinctions 

in macrofauna assemblages based on depth but found very distinct breaks correlated with sediment 

characteristics (Henkel and Politano 2017, Henkel and Nelson 2018). In this current study, spanning a 

much broader depth range, we did find major differences in macrofauna assemblages based on depth with 

subsequent distinctions related to sediment characteristics within major depth zones. This pattern follows 

that observed by Lissner (1989) in the northern and central California planning regions, where the major 

pattern in benthic community sampled between 90 and 600 m deep was related first to depth and then 

sediment-size characteristics, while other inter-basin differences (e.g., temperature and dissolved oxygen) 

appeared to have minor influence. Similarly, Bergen et al. (2001) sampled fauna from 10 to 200 m deep 

in southern California and found depth and secondary grain size-related distinctions, but did not identify 

any influence of latitude in defining assemblages. Our BEST result (Rho = 0.643 using just depth and 

median grain size) is numerically similar to the findings of Shumchenia and King (2010) from 

Narragansett Bay, where they found Rho = 0.689 with water depth, percent sand, and standard deviation 

of backscatter as variables.
21

We expected the shelf-slope break to be the most dramatic break in macrofauna assemblages. In Oregon 

the outer edge of the shelf has been reported as ranging 130 to 183 m deep (Byrne and Panshin 1977), and 

BOEM defines the edge of the OCS as 200 m consistent with Mendal (1964) and Davis (1972). However, 

the break in physical parameters resulting in the most distinctive assemblages was depth at ~90 m. 

Although unexpected, a transition at this depth has been reported before. While Bergen et al. (2001) 

classified the outer shelf as beyond 115 m, they explained that the transition between the two assemblages 

was found at depths of 90 to 115 m and cited earlier surveys conducted by the Hancock Foundation that 

found the deeper limit of their mid-depth assemblage to be 92 m (Barnard and Hartman 1959, Barnard 

and Ziesenhenne 1960, Jones 1964, 1969). This assemblage break at 90 m was also found when Henkel et 

al. (2014) used cluster analysis on the 2003 WEMAP macrofaunal dataset in the BOEM-BHC project. In

describing the distribution of organic carbon in surface sediment in the northeast Pacific Ocean, Gross et 

al. (1972) referred to the inner continental shelf as < 90 m and the outer continental shelf as 90 – 180 m, 

supporting the 90 m delineation in macrofaunal clusters found by our study; associating TOC differences 

with this boundary may suggest that sediment organic carbon may be the causative factor in this observed 

depth break on the shelf. While depth and TOC were strongly positively correlated at our sampled 

stations (Figure A-1) without an obvious break at 90 m (Figure A-2), deeper than 90 m, there are no 

stations with less than 0.15% TOC. In our current study, deeper than 90 m the next major break was at

200 m, consistent with what is considered the edge of the continental shelf (Mendal 1964, Davis 1972) 

and the major break in the dendrogram constructed by Lissner (1989) for central and northern California 

OCS macrofauna collected from 90 to 600 m deep.

Beyond 200 m, stations deeper than 445 m held distinct assemblages, a depth break similar to the edge of 

the mid-slope as reported by Lissner (450 m) and consistent with the break in Hyland’s (1991) stations 

where a break was detected between 410 and 565 m (no stations in between were sampled). Shallower 

than 90 m, we detected a depth break at ~43 m. This is slightly shallower than the distinct break in 

bivalve assemblages in Oregon reported by Voohries et al. (2018; 50 m), but the same as the deeper 

bounds of the shallow clusters determined in Bergen et al. (2001) for southern California. The approach 

used by Lissner (1989) and Bergen et al. (2001) was to build cluster dendrograms based on the 

macrofaunal assemblages, correlate those clusters with physical parameters, and then rank which 

parameters most contributed. In our approach, we used the LINKTREE routine to find breaks in the 

physical parameters that maximized differences among the biological assemblages at the stations. 

Considering that the studies were conducted in three different regions with overlapping but not 

continuous depth ranges using different statistical approaches, it is remarkable that similar depth 

delineations were found among them.

In addition to the species composition changes with depth, species richness also varied with depth. The 

number of species per grab (sample or areal richness) and the number of organisms per grab (abundance) 

declined with depth, and both of these trends also were described by Lissner (1989) and Oliver et al. 

(2011). Since the number of species can be positively related to the number of individuals, there is an 

argument that a decline in the number of species per area may simply be the result of having fewer

individuals in the sample (Gotelli and Colwell 2001). Standardizing by number of individuals avoids this 

statistical problem; although Oliver et al. (2011) argue for the importance of comparing communities in a 

known spatial context. In our study, when the number of species per grab was standardized by the number 

of organisms per grab the resulting numerical richness actually was highest at the deepest depth. At 

stations less than 90 m the numerical richness averaged 0.26, on the outer shelf (90 to 200 m) it averaged 

0.35, the Slope Break averaged 0.38, the Upper Slope averaged 0.39, and on the Slope (445 to 525 m) it 

was 0.59 as has been found in other studies where numerical species richness peaks along the slope (Rex 

1981, 1983; Levin et al. 2001). In contrast, Oliver et al. (2011) found that numerical species richness was 

quite similar across depths off California while Menot et al. (2010) found it to be highest along 

continental margins.
22

Within each of the depth zones, while further distinctions among macrofaunal assemblages were related 

to sediment characteristics – primarily median grain size – the biologically relevant breaks in grain size 

differed between the mid-shelf, outer-shelf, and slope. This lack of a consistent pattern across depths may 

be why other authors have reported grain size characteristics to be relatively more important at the local 

scale (e.g., Reiss et al. 2010). On the mid-shelf (43 – 90 m) in Oregon/Washington there appear to be 

three biologically defined habitats with increasing sediment grain size (III: < 223 μm, IV: <301 μm, and 

V: >315 μm). The </>223 μm break described herein is essentially the >/< 1% fine sediment described in 

Henkel and Politano (2017), and reinforces the additional break around 300 μm; it is not surprising as the 

137 stations sampled in that study were a subset of the 404 analyzed in this effort. However, this > 223 μm

(< 1% fines) group now includes stations from 45° to 46 °N (north of Newport and south of Nehalem, 

Oregon), which was a major latitudinal gap in Henkel and Politano (2017). These clean sand groups are 

notable due to the overall low species density (average ~ 23 taxa per grab) relative to other shelf groups 

(~37 taxa per grab).

In the silt-containing mid-shelf Oregon/Washington stations (< 223 μm), we detected a latitudinal gap 

between 43.8° and 44.5°N with groups of stations clustering north and south of that gap. This gap is, in 

part, due to the fact that no silt-containing stations were sampled on the mid-shelf of the central Oregon 

coast as the clean sand extends further offshore in this region. While these groups were delineated in the 

LINKTREE using latitude, they also represent a divergence of sediment types, with the northern stations 

averaging 189 μm (0.24% TOC) and the southern stations averaging 165 μm (0.35% TOC), although 

latitude was used because a clear threshold in grain size or TOC could not be correlated with the station 

groupings by the LINKTREE routine. Sediment grain size data are often correlated with infaunal 

community parameters to try to establish relationships between them (Henkel and Politano 2017). 

However, Snelgrove and Butman (1994) found little evidence that grain size alone was a causative factor 

in determining species distributions, as hydrodynamic energy and organic content of the sediment vary 

with grainsize and can directly influence feeding and settling dynamics of benthic fauna more so than the 

particle size. The mid-shelf, muddy northern California stations had ten times the percentage of fine 

sediment, much smaller grain size, and twice the TOC content compared to the stations at the same depth 

in the smallest grain size in Oregon. Although our current study did not resolve differences among 

sediment and other potential factors related to latitude, Henkel and Nelson (2018) also identified a 

latitudinal break in macrofaunal composition at the OR-CA border even after accounting for differences 

related to the >/< 5% fines threshold, indicating factors related to hydroclimate and/or species dispersal 

likely play a role.

On the outer shelf (90 – 200 m deep), differences in macrofaunal assemblages also distinguished among 

three grain size classes with breaks at ~110 μm and ~170 μm median grain size. Henkel and Politano 

(2018) reported a similar break at </>112 μm. When considering outer shelf stations with smaller 

grainsize (< 104 μm; groups VI and VII), northern and southern (break at 43.6°) assemblages were again 

distinguished as on the mid-shelf with the more northern group having slightly larger median grain size; 

however, on the outer shelf the northern stations also had higher percent TOC (1.38% vs. 0.91%). While 

this seems contrary to the classic relationship between grain size or percent fines and TOC, Gross et al. 

(1972) note that “although there is substantial variation, the amount of organic carbon in the sediment at 

any depth is usually...lowest near the OR-CA boundary”, indicating a latitudinal trend in TOC that may 

be driven by other factors. The Outer Shelf <104 μm South group (IV) also differed from the rest of the 

outer shelf groups by having warmer, more oxygenated bottom water. Finally, it contained a subsequent 

split between southern Oregon and stations from the northern San Andreas Fault area of California. This 

is consistent with the split on the mid-shelf separating our Eureka stations from southern Oregon and both 

are consistent with the break detected by Henkel and Nelson (2018) at the OR-CA border in the 2003 

EMAP samples. The Outer Shelf 111 – 167 μm group (VIII) was similar in average depth to the smaller 
23

grain size groups, but had approximately half the TOC content). The largest grain size group on the Outer 

Shelf (>170 um; IX) again had less than half the TOC content of group VIII.

On the slope, we identified three major depth ranges (as on the shelf) which we termed Slope Break, 

Upper Slope, and Slope. Further distinctions within those depth zones on the slope were not defined by 

the LINKTREE by breaks in sediment grain size; however, in the Upper Slope group (XI) a similar 

latitudinal break (at 44 °N) was detected as on mid-shelf. This latitudinal break corresponds to a break in 

sediment type, where more northern stations average larger grain size while the more southern stations 

average smaller; however, TOC conditions were similar in both regions at these Upper Slope stations. 

Among all the slope stations, grain sizes, percent fines, and TOC all were similar to the smallest grain 

size class defined on the outer shelf. However, percent fines and TOC increased with slope depth along 

with median grain size, temperature, oxygen, and pH decreasing, making it challenging to determine the 

actual drivers for assemblage differences across slope depths. Hyland et al. (1991) attributed depth-related 

patterns in abundances and diversity to a dissolved oxygen gradient. While we were not able to include 

dissolved oxygen as a potential factor in the BEST analysis (as we did not have data for all stations), it 

was the second highest correlate with MDS2 and is anti-correlated with depth.

3.4.1 Relationship to Offshore Development

Newer approaches are needed for sea floor planning and predicting an/or assessing impacts occurring at 

wide spatial scales. Macrofauna are an established tool to quantify the relative degree of anthropogenic 

impact, used most often in point source discharges occurring in defined areas of the ocean floor (Muxika 

et al. 2007, Ranasinghe et al. 2012). However, accurately assessing biotic condition over large spatial 

scales can be logistically challenging, especially when factoring in the water depths of the continental 

shelf and slope. Habitat mapping classified by surficial geology – used as a proxy for biological habitats –

has commonly been used in ocean sea floor planning decisions, yet many suggest changes to the sediment 

classifications within EUNIS (Galparsoro et al. 2012), and CMECs (Henkel and Politano 2017) need to 

be made to better reflect the biological communities (Shumchenia and King 2010).

An improved approach would be to quantify the relationship between the species distributions and the

environment, using local, faunal-based survey data to inform large-scale modeling of species occurrence

predicted from more easily measurable environmental data (Dutertre et al. 2013). Here, we used 

differences in species assemblages to find meaningful breaks in more easily measured environmental data 

in order to develop habitat classifications. For planning, these results can be used to set expectations of 

what macrofaunal species are likely to occur in particular habitat types. As a case study, we can apply our

soft bottom habitat characterizations to two marine hydrokinetic projects proposed in the PNW that would 

be expected to have direct or indirect impacts on soft sediment benthic communities (Dannheim et al.

2019; Boehlert and Gill 2010). The two projects are at different stages and follow distinct permitting 

pathways. Yet the results of this study can guide the design and define the reference areas and area of 

potential impact for both.

While both Oregon and northern California have strong wind and wave resources, are of interest for 

marine renewable energy development, and may have similar impacting factors to the seafloor, projects 

could be located in different sea floor soft bottom habitats with very different macrofaunal assemblages. 

PacWave South is in the final permitting stages offshore Newport, Oregon (central Oregon), to be a 

testing center for wave-powered marine hydrokinetic devices. The seafloor at PacWave South is Mid 

Shelf Clean Sand (Groups IV and V, average depth 63 m), which is a distinct medium to coarse sanddominated stretch of the Oregon central mid shelf. In contrast, offshore Eureka, California (northern 

California, average depth 900 m), where there is also interest in renewable energy, the seafloor is deeper. 

This area is deeper than the studied area, but is most closely associated with the deepest Slope habitat 

group (Group XII, average depth 484 m). That group has the highest TOC and percent fines of the areas 

sampled in this project, while also being in the latitudinal range of the Mid Shelf California Mud group 
24

(Group I, average depth 65 m), which has the second highest percent fines; thus we expect the area 

offshore Eureka to have high percentages of silt and TOC. In addition to hosting different macrofaunal 

assemblages, soft bottom community recovery differs depending on the local sediment and flow 

conditions (Kaiser et al. 2000, Bergman et al. 2015, Silberberger et al. 2019). We hypothesize then that 

the physical and biological responses to impacting factors and ecological recovery will be different in 

central mid shelf Oregon versus the upper slope of northern California. Having this expectation prior to 

development, instead of a post-hoc discovery, will aid in planning and the development of monitoring 

plans.

A regional characterization of the macrofaunal communities is needed to establish context for projects 

prior to development. However, the long-term need is to understand the alterations to ecological 

functioning and ecological processes that result from seafloor impacts (Dannheim et al. 2019). This 

means that to understand the macrofauna distribution at a finer temporal and spatial scale within a region, 

more hydrodynamic parameters need to be measured consistently (Dutertre et al. 2013). For artificial reef 

effects associated with offshore structures, TOC and nitrogen may become important signals as there is 

some evidence that artificial (oyster shell) reefs increase TOC and TN content in surface sediments (Xu et 

al. 2014). Further, to understand the impacts of development and the condition of the seafloor over time, 

the macrofauna species and communities should be linked to benthic condition indices. A site-specific 

response relies on knowing the functional role of the species of interest. Linking macrofauna condition to 

the ecological processes (Frid 2011) means that we need to next understand the forage value (Bond et al. 

1999) and trophic links to groundfish species important in the PNW. Finally, ecological function needs to 

be evaluated in the context of many other stressors such as changing climate and fishing impacts (Cada et 

al. 2007, Birchenough et al. 2015, Dannheim et al. 2019). This work is a step towards utilizing predictive 

modeling and quantifying species abiotic relationships with the aim at identifying ecosystem 

consequences of changes to various environmental factors (Méléder et al. 2010; Dutertre et al. 2013).

3.4.2 Conclusions

This study classified the macrofaunal communities of the PNW continental shelf and upper slope soft

bottom into 12 habitats. These habitats represent key depth and sediment-related breaks in macrofaunal 

community composition. Our results are consistent with the pattern of depth-related macrofauna 

assemblages throughout the California Current, supported by several studies in different parts of the 

California Current spanning over 30 years. Sediment characteristics were important parameters not just on 

the inner to mid-shelf as determined in Henkel and Politano (2017), but also further across the shelf and 

down the slope. In classifying these habitats, we can create a context for describing the current

communities in the region for soft bottom; a region undergoing changes in ocean hydrology, changes in 

bottom trawling pressure, and of growing interest for energy development projects. Establishing a soft 

bottom characterization for the PNW offshore will aid in the increasingly standardized and quantitative 

assessments of impacts.
25

4 Data Gathering and Species Selection for Region-wide Habitat 

Suitability Modeling

4.1 Purpose

A prior regional analysis of benthic habitat distribution (referenced as Benthic Habitat Characterization 

BOEM-BHC, Henkel et al. 2014) was motivated by wave energy project proposals in northern California 

and Oregon. Since that analysis, there has been greater interest in siting marine renewable energy projects 

in deeper waters of the continental slope, as well as waters further south in central and southern 

California. In response to this increase in geographic scope, we aimed to expand the depth and latitudinal 

range of habitat suitability models, like those developed in the BHC project, by incorporating other 

survey programs in California as well as Oregon. These models will be designed and run by NCCOS

(they are not updates of the same BayesNet models from the 2014 BHC project) and will be described in 

a separate BOEM report that NCCOS will develop under Interagency Agreement Number M16PG00014.

4.2 Study Area

The data for this chapter spanned from the California-Mexico border to Vancouver, Washington

(Figure 7). Sample data are from stations 3 to 1023 m deep.

4.3 Approach

Additional BOEM-funded research by the Southern California Coastal Water Research Project 

(SCCWRP) offered the opportunity to incorporate the many years of macrofaunal samples collected as 

part of the Southern California Bight Regional Monitoring Program (Schiff et al. 2016) to potentially 

expand the domain of the habitat suitability models to the entire lower US west coast. However, this 

combination of the OSU samples (spanning Fort Bragg, California, to Grays Harbor, Washington) plus 

the SCCWRP data left a large portion of the California coast without observations (e.g., Monterey, San 

Francisco, Humboldt). To fill these spatial gaps, we sought data from previous sampling programs carried 

out by various entities in the central and northern California region (e.g., EPA-WEMAP, EPA-Army 

Corps dredge disposal sampling, NOAA, municipalities). Thus, we endeavored to obtain as many of those 

records as possible to gain more complete coverage of the lower states of the US west coast.
26

Figure 7. Station locations for the 14 datasets included for modeling.

Details of each dataset are included in Table 6. The base map shows the surficial sediment percent sand and the 

200 m depth contour.
27

4.4 Data Collection and Processing

The Henkel lab provided many of the northern region datasets including data from this BOEM-funded 

study and the BOEM-BHC project as well as data from in and around the PacWave facilities operated by 

OSU’s Pacific Marine Energy Center and from a project off Reedsport and Coos Bay, Oregon, funded by 

the Oregon Wave Energy Trust. The EMAP data are available for download from the National Coastal 

Assessment Coastal Data Search Engine: https://oaspub.epa.gov/coastal/coast.search. The 2003 EMAP 

data for California north of Point Conception, Oregon, and Washington were included in the MS Access 

database developed by the Henkel lab. Additional SCCWRP data was obtained directly from David 

Gillett and used for determining model species but were not included in the MS Access database. 

ODMDS data from Oregon was obtained directly from the US Army Corps of Engineers (USACE), 

facilitated by previous collaborative projects between Henkel and USACE/US Environmental Protection 

Agency (EPA). Metadata from the MCI, CLLEAN, and NOAA Monterey sampling are available on the 

MLML website and data were subsequently provided by MLML upon request. The SF Offshore and 

HOODS data were provided by Walt Nelson, former Asst. to Division Director of the Western Ecology 

Division of the EPA. Shallow water Morro Bay station data were obtained from the City of Morro Bay 

and Cayucos Sanitary District Offshore Monitoring and Reporting Program available from the city’s 

website: http://www.morro-bay.ca.us/ArchiveCenter/ViewFile/Item/2757. Collectively, these datasets 

spanned 1994 to 2016 from 15 to 535 m deep for a total of 1835 records (Table 5). All records contained 

macrofaunal count data, station location, depth, and at least percent fines for the sediment. Some datasets 

had additional physical data.

All data from north of Point Conception were entered into a Microsoft Access database and a "taxon 

reduced" query was created from the various data sets. Because projects often had different spelling for 

the same animal (e.g., Acteocina sp vs Acteocina spp vs Acteocina spp.) over 3000 unique taxa were 

generated. Sometimes different scientific names were used, including older names no longer recognized 

as valid by the scientific community. Additionally, there were many misspelled entries and common 

names were occasionally used. In some instances, the stage of the animal was included in the text (e.g.,

Aeolidiacea sp. Juv.) and there were other name differences making merging the data sets difficult. So, a 

secondary column was created and used in a query to aggregate all the similar species together. The 

World Register of marine species (WoRMS; accessed in 2017) was used to cross check species names 

and select valid species names. In some instances, it was not possible to determine what the animal was at 

the species level and a taxa name was chosen at the lowest possible level (e.g., Cuke-like Anemone 

became Actiniaria sp). It was from this query “Total Taxon Reduced” that all data were pulled for the 

statistical analyses described below.
28

Table 6. Datasets collated by Henkel lab and included in Microsoft Access Database.

Dataset 

label (Fig 7)

Collection 

type

Sieve mesh 

size (mm) Region Years

Depth 

range 

(m)

No. of 

Records

Dataset sources

BHC Box core 1 WA, OR, 

northern CA 2010-2012 49-133 150 Benthic habitat characterization (BHC) surveys from 

Oregon State University (OSU); Henkel et al. 2014

WEMAP van Veen 1 WA, OR, CA 2003 28-126 147

Environmental monitoring & assessment program 

(EMAP) data provided by Environmental Protection 

Agency (EPA); Nelson et al. 2008.

ODMDS Box core 0.5 OR

2008, 2009, 

2013, 2014, 

2016

10-85 309

Designated ocean dredged material disposal sites 

(ODMDS) from 7 Oregon sites provided by EPA 

Region 10

HOODS van Veen 0.5 northern CA 2008, 2014 31-92 44 Humboldt open ocean disposal site (HOODS) data 

provided by EPA Region 9

Farallones van Veen 1 central CA 2009 8-34 12

Assessing potential resource utilization by Gray 

Whales in the Gulf of the Farallones National Marine 

Sanctuary; data provided by EPA Region 9

SFPUC van Veen 0.5 central CA 2004, 2005, 

2010 12-35 154 San Francisco offshore provided by the San 

Francisco Public Utility Commission

MCI Smith-McIntyre 0.5 central CA 1999 10-450 83

Surveys in Monterey Bay National Marine Sanctuary 

for MCI Worldcom (ABA Consultants 2000) data 

provided by Moss Landing Marine Laboratories 

(MLML) Benthic Lab

CCLEAN Smith-McIntyre/ 

van Veen 0.5 central CA

2001-06, 

2008-10, 

2015

80 90

Central Coast Long-term Environmental 

Assessment Network (CCLEAN) data provided by 

MLML Benthic Lab

NOAAMLML Smith-McIntyre 0.5 central CA 2004, 2005 80-476 53 Data collected by NOAA and provided by MLML

Benthic Lab

Morro Bay Young Modified 

van Veen 1 central CA 2015 15 7 City of Morro Bay and Cayucos Sanitary District 

Offshore Monitoring and Reporting Program

PacWave Box core 1 central OR 2010-2016 20-70 597 PacWave test sites data collected by Henkel funded 

by various sources

OWET Box core 1 southern OR 2011 24-90 42 Data collected by Henkel funded by Oregon Wave 

Energy Trust (OWET)

OSU-BOEM Box core 1 OR 2014-2016 60-525 147 Current report
29

4.5 Statistical Analyses

We first attempted to conduct multivariate analyses on the entire dataset. These analyses included 

creating a dissimilarity matrix and producing non-parametric MDS plots and cluster dendrograms to 

visualize how stations grouped together based on the macrofaunal assemblages collected. Including all of 

the records from the southern California Bight made the dataset too big for PRIMER or R to handle in a 

timely manner. So, we first analyzed the data included in the MS Access database only north of Point 

Conception. Within this study area, multivariate analysis indicated a strong break between samples 

collected using a 1.0 mm mesh sieve and those collected using a 0.5 mm mesh. Thus, analyses to identify 

candidate species for modeling (not the final modeling) were conducted using only samples obtained by 

using a 1.0 mm mesh size. Once this approach was decided, we then used the Bight Survey data from just 

2003 at this stage as those were temporally consistent with the rest of the WEMAP samples, which was 

the major source of station data for central and northern California (Nelson et al. 2008). Using this final 

dataset for assemblage determination (n = 1483 stations), we conducted cluster analysis with SIMPROF 

set at the 1% significance level to determine statistically significant groups of the 1483 stations followed 

by SIMPER analyses to determine which species contributed to the similarities within groups and 

differences between them.

We then determined the number of times each species was indicated as being significant contributors to 

the similarities within groups overall and within each ecoregion (OR/WA, NorCal, Southern California 

Bight). Species were then considered for potential as model species if they were frequently significant 

contributors to the similarities within group, or if they were determined to be frequent distinguishers 

between the 12 major habitat groups in the in the previous chapter. These species were added separately 

because they included species that were discriminating on the slope, where we had little coverage in the 

remainder of the study region. Thus, in the analysis of the entire region, they were not necessarily 

frequently occurring characteristic species. Additional refinements were made for the final list of 

candidate species for modeling. For example, species that were significant contributors to the similarities 

within groups for only one ecoregion were only considered if we had additional evidence that their 

presence/absence contributed to differences among habitats within that region (and weren’t just defining 

the region). Because samples were biased to the northern part of the region, when one species in a genus 

was a frequent distinguisher in the north and a conspecific was more common in the south, we added the 

southern conspecific to the list of species to be modeled. We also prioritized species that had a p-code 

pollution tolerance scores of less than 30 in the SCCWRP database, favoring species that were 

representative of non-impacted conditions. (For the p-codes, a higher number indicates greater tolerance 

to pollution; Smith et al. 2001). Finally, if not already included, we added taxa that we had previously 

modeled in the BOEM-BHC project in order to compare model outputs between model approaches.

Finally, we checked the frequency of occurrence of each of the selected species to ensure there were 

enough occurrences (not just large abundances at a few stations) to be useful for modeling. In order to be 

considered for modeling, we aimed for at least 10% overall occurrence in the northern study region; 

however, this was not always possible for slope species as the number of slope samples was low relative 

to the entire dataset. While the initial lists of potential model species were derived based only on the 

1 mm mesh sieve studies, occurrence data was based on all the datasets.

4.6 Results

The cluster analysis with a SIMPROF threshold of 1% resulted in 303 significantly different groups. 

There were 147 significant groups detected in the PNW Region (Oregon and Washington), 29 groups in 

the Northern California (above Point Conception) Region, and 127 groups in Southern California (the 

Bight data). Forty-three taxa were selected as the final list recommended for future habitat suitability 

modeling (Table 7). Polychaetes were the most frequent group considered to be characteristic of and/or 
30

distinguishing between macrofaunal assembalges and 22 taxa were selected for modeling; taxonomic 

changes over the course of the sampling efforts resulted in some suspected synomyns being lumped 

together. Molluscs were the second most frequent characteristic/distinguishing taxa, and eight bivalves, 

six gastropods, and the caudofoveatan, Chaetoderma argenteum (which was a slope distinguishing 

species), were selected for modeling. The most frequently overall characteristic species (contributing to 

the similarity of the stations within a group) was the ubiquitous bivalve, Axinopsida serricata, which was 

characteristic in 95 of the 303 groups. Thus, while widespread, we would expect to model some 

unsuitable habitat for this species as it was not found to be highly characteristics of all macrofaunal 

assemblages. Only three crustacea were determined to be highly characteristic of assemblages or 

distinguishing among them: the mud-tolerant shelf amphipod, Ampelisca careyi, the highly abundant pea 

crab, Pinnixa occidentalis complex, and the ostracod, Euphilomedes carcharodonta. Three echinoderms 

were also determined to be highly characteristic of assemblages or distinguishing among them: two 

ophiuroids and the heart urchin, Brisaster latifrons, which was indicative of the outer shelf and slope with 

the smallest grain sizes.

The results of the models (maps, envrionmental predictors, and model summaries) will be provided and 

discussed in a subsequent publication from NCCOS.
31

Table 7. The 43 taxa recommended for future habitat suitability modeling.

The Times Characteristic Total column (Times Char Total) indicates the number of times a taxon was characteristic in statistically significant clustered groups

(n = 303 total clusters). The percentage of groups for which a taxon was characteristic are separated into three latitudinal areas from north to south: PNW (Oregon 

and Washington), NorCal (California, north of Point Conception), and Bight (California, south of Point Conception, Southern California Bight). The North Occur 

column indicates the number of stations at which taxa were collected north of Point Conception (PWN and NorCal combined). The Bight Occur column indicates 

the number of times taxa were collected during sampling in the Southern California Bight (Gillet et al. 2017). The total possible number of occurrences in the north 

is 1795 (the total number of sampling events or station visits) and 1209 in the Southern California Bight. Bight p-code values are the relative degree of pollution

tolerance/sensitivity quantified by taxon and by depth in the Southern California Bight (Smith et al. 2001). The plus symbol (

+) indicates species discriminating 

among habitats in the previous chapter. The asterisk symbol (*) indicates the seven species that were modeled for the BOEM-BHC project (Henkel et al. 2014).

Times 

Char 

Total

% of 

Groups 

PNW

% of 

Groups 

NorCal

% of 

Groups 

Bight

North 

Occur

Bight 

Occur

Bight

p-code

(0-30 m)

Bight

p-code

(30-120 m)

Bight

p-code

(120-300 m)

Acila castrensis 29 18% 7% 1% 302 33 -10.711 25.26

Adontorhina cyclia+ 30 16% 10% 3% 171 107 -21.116 -7.526

Alia (Astyris) gausapata* 28 19% 562 1 No p-code

Ampelisca careyi 57 22% 28% 13% 597 426 16.113 -11.303 -7.789

Amphiodia urtica 51 12% 24% 21% 452 133 48.713 -8.565 -12.225

Amphioplus macraspis+ 11 7% 84 0 Not present

Axinopsida serricata* 95 48% 34% 12% 936 367 69.746 26.965 60.36

Brisaster latifrons+ 18 9% 3% 3% 158 0 Brisaster sp. 2.95

Callianax pycna* 18 12% 423 1 No p-code

Chaetoderma argenteum+ 5 3% 62 0 Not present; other Chaetoderma spp listed

Chloeia pinnata+ 31 3% 10% 19% 179 400 37.607 19.019 26.917

Cylichna attonsa 53 36% 812 0 Not present; other Cylichna spp listed

Ennucula tenuis* 34 18% 10% 4% 412 141 15.558 -0.374 -4.549

Euphilomedes carcharodonta 52 12% 24% 21% 368 432 71.111 59.539 42.635

Gadila tolmiei+ 7 5% 59 35 No p-code

Galathowenia oculata+ 25 15% 10% 323 4 -2.075 -3.221 -2.633

Glycera nana/tesselata 44 11% 28% 16% 416 509 50.612 39.829 53.637

Glycinde armigera+ 52 22% 17% 12% 750 372 19.129 19.126 38.526

Huxleyia munita+ 18 12% 1% 126 30 -66.319
32

Times 

Char 

Total

% of 

Groups 

PNW

% of 

Groups 

NorCal

% of 

Groups 

Bight

North 

Occur

Bight 

Occur

Bight

p-code

(0-30 m)

Bight

p-code

(30-120 m)

Bight

p-code

(120-300 m)

Kurtiella tumida 42 11% 31% 13% 585 361 45.575 59.199 51.633

Leitoscoloplos pugettensis+ 51 20% 17% 600 206 42.489 47.53 8.019

Macoma carlottensis 35 22% 10% 441 87 106.04 115.822 76.711

Magelona berkeleyi* 4 3% 125 66 26.808 29.525 -29.055

Magelona sacculata 40 27% 3% 895 25 -8.299 32.344

Maldane sarsi+ 27 7% 24% 7% 312 364 18.308 9.273 17.309

Ninoe gemmea 12 5% 14% 127 0 42.018 23.234 41.577

Ninoe tridentata 3 3.4% 1.6% 76 205 42.018 23.234 41.577

Nutricola lordi+ 27 18% 305 3 No p-code

Onuphis iridescens* 55 30% 24% 3% 755 140 19.245 29.726 35.694

Paraprionospio alata/pinnata 60 12% 31% 26% 498 795 10.608 21.746 38.608

Phoronidae sp+ 36 11% 7% 14% 75 449 17.859 8.077 -2.306

Pinnixa occidentalis complex 29 11% 31% 3% 292 29 39.228 24.216 41.247

Polycirrus spp 8 7% 5% 246 439 -1.05 -5.026 1.856

Praxillella gracilis+ 3 1% 3% 98 33 12.968 15.645 10.416

Prionospio (prionospio) jubata 42 6% 21% 21% 159 596 55.502 31.466 32.647

Pulsellum salishorum+ 11 7% 135 0 Not present

Rhabdus rectius+ 23 14% 7% 210 58 34.765 41.919 29.978

Scoletoma luti 24 12% 21% 455 0 49.381 30.228 19.076

Spiophanes berkeleyorum/kimballi 62 16% 38% 22% 734 692 24.199 33.753 38.763

Spiophanes duplex 54 14% 39% 0 800 6.131 8.546 -1.569

Spiophanes norrisi 60 29% 14% 10% 102 353 -2.309 12.055 -23.746

Sternaspis assimilis+ 3 2% 39 0 Not present; other Sternaspis spp listed

Sternaspis affinis/fossor* 37 9% 24% 13% 283 405 34.246 -17.301 -1.458
33

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38

Appendix: Additional Tables and Figures in Support of Chapter 3

Table A-1. Station data for the 404 samples considered in Chapter 3.

Station Year Latitude Longitude Depth

42.0_060m2014 2014 42.0035 -124.366 56

42.0_060m2015 2015 42.00153 -124.373 64

42.0_100m2014 2014 42.00345 -124.471 99

42.0_100m2015 2015 42.01683 -124.470 103

42.0_200m2014 2014 41.99835 -124.589 215

42.0_200m2015 2015 42.00127 -124.586 202

42.0_300m2014 2014 41.9948 -124.605 329

42.0_300m2015 2015 42.00163 -124.602 306

42.0_400m2014 2014 41.99827 -124.617 394

42.0_400m2015 2015 42.00193 -124.618 400

42.5_100m2014 2014 42.50048 -124.657 100

42.5_200m2015 2015 42.50198 -124.731 216

42.5_300m2014 2014 42.50257 -124.749 308

42.5_300m2015 2015 42.50048 -124.756 302

42.5_400m2014 2014 42.50205 -124.768 400

42.5_400m2015 2015 42.50128 -124.770 396

43.0_060m2014 2014 43.00450 -124.521 59

43.0_100m2014 2014 43.00667 -124.594 99

43.0_100m2015 2015 42.99173 -124.628 112

43.0_200m2014 2014 42.99922 -124.854 202

43.0_300m2014 2014 43.00203 -124.869 288

43.0_300m2015 2015 43.00000 -124.870 295

43.0_400m2014 2014 43.00275 -124.899 398

43.0_400m2015 2015 42.99938 -124.898 395

43.5_060m2014 2014 43.51248 -124.297 61

43.5_060m2015 2015 43.50038 -124.302 63

43.5_100m2014 2014 43.49880 -124.381 101

43.5_100m2015 2015 43.49542 -124.413 110

43.5_200m2014 2014 43.49990 -124.599 204

43.5_200m2015 2015 43.49928 -124.598 206

43.5_300m2014 2014 43.50262 -124.644 299

43.5_300m2015 2015 43.49900 -124.644 300

43.5_400m2014 2014 43.50285 -124.678 399

43.5_400m2015 2015 43.49900 -124.677 401

43.5_500m2014 2014 43.50272 -124.720 497

43.5_500m2015 2015 43.50017 -124.721 507

44.0_150m2014 2014 43.99732 -124.569 148

44.0_150m2015 2015 43.99983 -124.581 155
39

Station Year Latitude Longitude Depth

44.0_200m2014 2014 44.00373 -124.940 217

44.0_200m2015 2015 43.99917 -124.940 209

44.0_300m2014 2014 44.00635 -124.953 309

44.0_300m2015 2015 43.99815 -124.955 306

44.0_400m2014 2014 44.00435 -124.957 352

44.0_400m2015 2015 43.99680 -124.960 393

44.5_100m2015 2015 44.49817 -124.532 107

44.5_200m2015 2015 44.49983 -124.688 202

44.5_300m2015 2015 44.49983 -124.746 307

44.5_400m2014 2014 44.50278 -124.884 359

44.5_400m2015 2015 44.50333 -124.792 394

45.0_060m2014 2014 44.99967 -124.090 61

45.0_060m2015 2015 44.9967 -124.090 65

45.0_100m2014 2014 44.99977 -124.164 101

45.0_100m2015 2015 44.99917 -124.165 107

45.0_200m2014 2014 45.00467 -124.359 200

45.0_200m2015 2015 44.99983 -124.359 204

45.0_300m2014 2014 45.00338 -124.439 300

45.0_300m2015 2015 45.00067 -124.438 304

45.0_400m2014 2014 45.00492 -124.521 402

45.0_400m2015 2015 44.99983 -124.517 407

45.5_060m2014 2014 45.49530 -124.088 66

45.5_060m2015 2015 45.51250 -124.053 64

45.5_100m2014 2014 45.50303 -124.150 107

45.5_100m2015 2015 45.50583 -124.133 104

45.5_200m2014 2014 45.49460 -124.424 208

45.5_200m2015 2015 45.49600 -124.414 207

45.5_300m2014 2014 45.48427 -124.469 294

45.5_300m2015 2015 45.48850 -124.471 306

45.5_400m2014 2014 45.48588 -124.502 408

45.5_400m2015 2015 45.48100 -124.497 405

46.0_060m2015 2015 46.00150 -124.066 61

46.0_100m2015 2015 46.00000 -124.239 106

46.0_200m2014 2014 46.00088 -124.673 201

46.0_200m2015 2015 45.99950 -124.678 206

46.0_300m2014 2014 45.98730 -124.722 300

46.0_300m2015 2015 46.00000 -124.728 296

46.0_400m2014 2014 45.98842 -124.762 409

46.0_400m2015 2015 45.99917 -124.770 405

BA543_Sept12 2012 43.04990 -124.620 128

BA544_Sept12 2012 43.0331 -124.584 100
40

Station Year Latitude Longitude Depth

BA548_Aug12 2012 43.0657 -124.568 91

BA551_Sept12 2012 43.0502 -124.604 128

BA552_Aug12 2012 43.1016 -124.602 128

BA555_Sept12 2012 43.0486 -124.533 91

BA556_Aug12 2012 43.1007 -124.551 91

BA560_Sept12 2012 43.0674 -124.585 109

BA563_Sept12 2012 43.0333 -124.533 73

BA564_Aug12 2012 43.0827 -124.534 73

BA568_Aug12 2012 43.0998 -124.584 100

BA572_Aug12 2012 43.0664 -124.534 109

BA576_Sept12 2012 43.0659 -124.587 118

BA579_Sept12 2012 43.0495 -124.568 100

BA580_Aug12 2012 43.0994 -124.539 67

BB20A_Jun11 2011 44.726 -124.090 21.9

BB20B_Jun11 2011 44.72633 -124.090 21.1

BB30A_Jun11 2011 44.72633 -124.100 30.2

BB30B_Jun11 2011 44.72633 -124.100 29.9

BB40A_Jun11 2011 44.72467 -124.110 42.5

BB40B_Jun11 2011 44.72467 -124.110 42

BB50A_Jun11 2011 44.72367 -124.120 49.7

BB50B_Jun11 2011 44.72333 -124.120 49.5

BB60A_Jun11 2011 44.7275 -124.150 59.7

BB60B_Jun11 2011 44.72667 -124.150 59.6

BB70A_Jun11 2011 44.71767 -124.210 72.5

BB70B_Jun11 2011 44.72217 -124.210 72.1

CABLE12016 2016 43.42262 -124.637 193

CABLE22016 2016 43.41898 -124.578 144

CABLE32016 2016 43.42395 -124.519 114

CABLE42016 2016 43.42493 -124.459 104

CABLE52016 2016 43.41992 -124.415 89

CABLE62016 2016 43.42522 -124.341 34

CABLE72016 2016 43.42648 -124.311 20

CABLE82016 2016 43.42322 -124.683 338

CP541_Oct12 2012 44.283 -124.248 70

CP542_Oct12 2012 44.2355 -124.269 74

CP545_Oct12 2012 44.2583 -124.234 66

CP546_Oct12 2012 44.2287 -124.242 69

CP549_Oct12 2012 44.2943 -124.216 61

CP550_Oct12 2012 44.2473 -124.284 75

CP553_Oct12 2012 44.2412 -124.220 62

CP554_Oct12 2012 44.2867 -124.272 73
41

Station Year Latitude Longitude Depth

CP557_Oct12 2012 44.2735 -124.196 56

CP558_Oct12 2012 44.222 -124.292 77

CP561_Oct12 2012 44.2112 -124.216 62

CP562_Oct12 2012 44.2823 -124.309 78

CP565_Oct12 2012 44.2792 -124.250 69

CP566_Oct12 2012 44.2182 -124.252 72

CP569_Oct12 2012 44.209 -124.248 71

CP570_Oct12 2012 44.2058 -124.294 78

CP573_Oct12 2012 44.263 -124.223 62

CP574_Oct12 2012 44.2143 -124.305 80

CP577_Oct12 2012 44.2368 -124.203 58

CP578_Oct12 2012 44.2573 -124.297 77

EUR002_Oct10 2010 40.914 -124.222 52

EUR006_Oct10 2010 40.8789 -124.260 61

EUR018_Oct10 2010 40.9021 -124.236 55

EUR022_Oct10 2010 40.8593 -124.254 55

EUR034_Oct10 2010 40.9139 -124.283 75

EUR038_Oct10 2010 40.8656 -124.374 123

EUR045_Oct10 2010 40.8841 -124.292 73

EUR050_Oct10 2010 40.9204 -124.314 89

EUR054_Oct10 2010 40.9144 -124.337 102

EUR061_Oct10 2010 40.8676 -124.309 75

EUR066_Oct10 2010 40.9086 -124.261 64

EUR074_Oct10 2010 40.9197 -124.289 77

EUR082_Oct10 2010 40.9116 -124.330 97

EUR090_Oct10 2010 40.8728 -124.253 57

EUR098_Oct10 2010 40.8759 -124.354 104

EUR101_Oct10 2010 40.8578 -124.295 68

EUR106_Oct10 2010 40.8432 -124.339 79

EUR109_Oct10 2010 40.8862 -124.285 71

EUR114_Oct10 2010 40.9159 -124.354 118

EUR117_Oct10 2010 40.8429 -124.310 57

GH004_Sept10 2010 46.9312 -124.393 58

GH020_Sept10 2010 46.9783 -124.397 56

GH047_Sept10 2010 46.9468 -124.502 82

GH063_Sept10 2010 46.9296 -124.481 78

GH076_Sept10 2010 46.9781 -124.449 66

GH084_Sept10 2010 46.9688 -124.404 60

GH092_Sept10 2010 46.9815 -124.423 63

GH103_Sept10 2010 46.9758 -124.465 71

GH111_Sept10 2010 46.9571 -124.500 80
42

Station Year Latitude Longitude Depth

GH119_Sept10 2010 46.9783 -124.509 79

GH122_Sept10 2010 46.9619 -124.463 71

GH165_Sept10 2010 46.956 -124.420 63

GH221_Sept10 2010 46.9783 -124.418 61

MB20A_Jun11 2011 44.69567 -124.090 19.9

MB20B_Jun11 2011 44.69567 -124.090 19.6

MB30A_Jun11 2011 44.691 -124.100 30.1

MB30B_Jun11 2011 44.69383 -124.100 30.2

MB40A_Jun11 2011 44.6925 -124.110 41.7

MB40B_Jun11 2011 44.6925 -124.110 40.6

MB50A_Jun11 2011 44.692 -124.140 52.6

MB50B_Jun11 2011 44.69183 -124.140 52.2

MB60A_Jun11 2011 44.6905 -124.150 59.6

MB60B_Jun11 2011 44.52267 -124.160 59.4

MB70A_Jun11 2011 44.69033 -124.220 69.6

MB70B_Jun11 2011 44.68967 -124.230 69.6

MC102016 2016 43.74947 -124.621 307

MC112016 2016 43.74995 -124.700 459

MC12016 2016 43.55032 -124.568 195

MC122016 2016 43.900 -124.550 162

MC132016 2016 43.89965 -124.600 215

MC142016 2016 43.89992 -124.699 224

MC152016 2016 44.05037 -124.550 132

MC162016 2016 44.05015 -124.600 138

MC172016 2016 44.12985 -124.720 112

MC182016 2016 44.13042 -124.550 114

MC192016 2016 44.12983 -124.589 123

MC202016 2016 44.05 -124.70 126

MC212016 2016 44.1984 -124.551 105

MC22016 2016 43.5507 -124.706 500

MC222016 2016 44.20003 -124.598 110

MC232016 2016 44.19985 -124.654 114

MC242016 2016 44.37492 -124.353 75

MC272016 2016 44.82057 -124.149 64

MC282016 2016 44.82028 -124.272 116

MC312016 2016 44.9203 -124.463 252

MC32016 2016 43.54563 -124.652 363

MC332016 2016 45.02668 -124.146 102

MC342016 2016 45.00078 -124.360 201

MC362016 2016 45.09857 -124.121 92

MC372016 2016 45.09978 -124.350 240
43

Station Year Latitude Longitude Depth

MC382016 2016 45.0999 -124.450 352

MC392016 2016 45.19963 -124.195 151

MC402016 2016 45.20003 -124.270 180

MC412016 2016 45.1996 -124.352 330

MC42016 2016 43.64325 -124.551 195

MC422016 2016 45.19977 -124.452 405

MC432016 2016 45.2996 -124.179 132

MC442016 2016 45.30025 -124.280 186

MC452016 2016 45.30383 -124.458 265

MC462016 2016 45.29953 -124.455 400

MC472016 2016 45.39643 -124.106 93

MC482016 2016 45.40038 -124.249 162

MC492016 2016 45.40818 -124.405 245

MC502016 2016 45.39978 -124.453 377

MC52016 2016 43.64968 -124.520 162

MC522016 2016 45.48963 -124.353 183

MC62016 2016 43.649 -124.700 493

MC72016 2016 43.65158 -124.648 375

MC82016 2016 43.75062 -124.550 185

MC92016 2016 43.74967 -124.600 260

NEH007_Sept10 2010 45.7112 -124.109 87

NEH011_Sept10 2010 45.6586 -124.156 104

NEH015_Sept10 2010 45.6552 -124.033 63

NEH023_Sept10 2010 45.6713 -124.140 99

NEH027_Sept10 2010 45.653 -124.053 70

NEH031_Sept10 2010 45.7269 -124.068 76

NEH035_Sept10 2010 45.6469 -124.147 101

NEH039_Sept10 2010 45.6702 -124.085 81

NEH043_Sept10 2010 45.6924 -124.049 71

NEH051_Sept10 2010 45.7264 -124.135 94

NEH055_Sept10 2010 45.7094 -124.130 94

NEH059_Sept10 2010 45.705 -124.027 64

NEH071_Sept10 2010 45.6552 -124.070 76

NEH075_Sept10 2010 45.646 -124.061 73

NEH079_Sept10 2010 45.6997 -124.043 69

NEH083_Sept10 2010 45.7208 -124.109 86

NEH087_Sept10 2010 45.6749 -124.036 67

NEH091_Sept10 2010 45.6908 -124.102 86

NEH095_Sept10 2010 45.7229 -124.039 62

NEH099_Sept10 2010 45.6625 -124.122 94

NEH107_Sept10 2010 45.6968 -124.114 88
44

Station Year Latitude Longitude Depth

NEH115_Sept10 2010 45.7288 -124.153 100

NH20A_Jun11 2011 44.6535 -124.090 22.4

NH20B_Jun11 2011 44.65333 -124.090 21.6

NH30A_Jun11 2011 44.6535 -124.100 32.6

NH30B_Jun11 2011 44.6535 -124.100 32

NH40A_Jun11 2011 44.652 -124.110 40.9

NH40B_Jun11 2011 44.652 -124.110 41.6

NH50A_Jun11 2011 44.65283 -124.150 49.3

NH50B_Jun11 2011 44.6525 -124.150 49.2

NH60A_Jun11 2011 44.65383 -124.180 58.9

NH60B_Jun11 2011 44.65317 -124.180 58.8

NH70A_Jun11 2011 44.65167 -124.230 71

NH70B_Jun11 2011 44.65117 -124.230 70.9

NPT003_Oct10 2010 44.6545 -124.243 74

NPT010_Oct10 2010 44.687 -124.197 69

NPT013_Oct10 2010 44.6323 -124.225 67

NPT014_Oct10 2010 44.704 -124.197 67

NPT019_Oct10 2010 44.6375 -124.271 77

NPT026_Oct10 2010 44.67 -124.192 67

NPT030_Oct10 2010 44.718 -124.158 61

NPT042_Oct10 2010 44.6838 -124.152 55

NPT046_Oct10 2010 44.6553 -124.226 70

NPT057_Oct10 2010 44.626 -124.148 50

NPT058_Oct10 2010 44.658 -124.188 62

NPT062_Oct10 2010 44.6888 -124.226 69

NPT067_Oct10 2010 44.7253 -124.223 73

NPT070_Oct10 2010 44.686 -124.161 58

NPT078_Oct10 2010 44.6472 -124.206 66

NPT085_Oct10 2010 44.6185 -124.190 57

NPT086_Oct10 2010 44.63 -124.150 49

NPT093_Oct10 2010 44.6265 -124.260 74

NPT094_Oct10 2010 44.7203 -124.206 70

NPT102_Oct10 2010 44.6585 -124.210 68

NPT110_Oct10 2010 44.6785 -124.243 72

NPT118_Oct10 2010 44.6745 -124.225 70

NS20A_Jun11 2011 44.62817 -124.090 18.4

NS20B_Jun11 2011 44.62817 -124.090 18.2

NS30A_Jun11 2011 44.62583 -124.110 35.3

NS30B_Jun11 2011 44.62583 -124.110 36.7

NS40A_Jun11 2011 44.6215 -124.120 40.3

NS40B_Jun11 2011 44.62133 -124.120 40.3
45

Station Year Latitude Longitude Depth

NS50A_Jun11 2011 44.6265 -124.130 48.6

NS50B_Jun11 2011 44.62633 -124.130 48.7

NS60A_Jun11 2011 44.62317 -124.190 57.7

NS60B_Jun11 2011 44.62067 -124.190 56.5

NS70A_Jun11 2011 44.624 -124.240 73.8

NS70B_Jun11 2011 44.622 -124.240 71.5

NSAF012_Oct10 2010 39.6979 -123.895 118

NSAF016_Oct10 2010 39.6441 -123.882 115

NSAF024_Oct10 2010 39.858 -124.335 122

NSAF028_Oct10 2010 39.8007 -123.920 102

NSAF032_Oct10 2010 39.5877 -123.864 107

NSAF040_Oct10 2010 39.842 -123.969 115

NSAF044_Oct10 2010 39.7632 -123.914 116

NSAF048_Oct10 2010 39.5948 -123.882 118

NSAF056_Oct10 2010 39.8499 -123.976 113

NSAF060_Oct10 2010 39.6791 -123.879 111

NSAF064_Oct10 2010 39.5177 -123.909 125

NSAF072_Oct10 2010 39.7913 -123.943 127

NSAF080_Oct10 2010 39.642 -123.855 95

NSAF088_Oct10 2010 39.6255 -123.889 120

NSAF096_Oct10 2010 39.6335 -123.905 127

NSAF100_Oct10 2010 39.8725 -124.071 122

NSAF104_Oct10 2010 39.6123 -123.883 98

NSAF108_Oct10 2010 39.8462 -123.996 122

NSAF112_Oct10 2010 39.6128 -123.899 125

NSAF116_Oct10 2010 39.718 -123.915 129

NSAF120_Oct10 2010 39.48 -123.923 118

PD30_Jun15 2015 44.591 -124.115 30.5

PD40_Jun15 2015 44.5923 -124.121 39.7

PD50_Jun15 2015 44.5937 -124.160 51.8

PD60_Jun15 2015 44.5937 -124.206 60.6

PD70_Jun15 2015 44.5902 -124.233 70.1

RP301_Jun11 2011 43.7325 -124.215 28.4

RP302_Jun11 2011 43.8182 -124.192 24.3

RP304_Jun11 2011 43.7663 -124.212 35.1

RP305_Jun11 2011 43.7883 -124.209 35.9

RP306_Jun11 2011 43.7407 -124.216 32.4

RP307_Jun11 2011 43.7352 -124.228 42.8

RP308_Jun11 2011 43.7913 -124.210 37.8

RP309_Jun11 2011 43.7625 -124.218 39.4

RP310_Jun11 2011 43.7965 -124.231 63.1
46

Station Year Latitude Longitude Depth

RP312_Jun11 2011 43.7445 -124.240 61.5

RP313_Jun11 2011 43.7834 -124.235 65.7

RP314_Jun11 2011 43.7408 -124.243 65

RP315_Jun11 2011 43.816 -124.229 64.1

RP316_Jun11 2011 43.7385 -124.251 72.7

RP317_Jun11 2011 43.766 -124.244 71.7

RP318_Jun11 2011 43.8003 -124.242 73.7

RP319_Jun11 2011 43.74 -124.255 79.5

RP320_Jun11 2011 43.7333 -124.257 79.2

RP321_Jun11 2011 43.7387 -124.255 77.9

RP326_Jun11 2011 43.7485 -124.214 32.1

RP336_Jun11 2011 43.741 -124.236 54.8

RS40_Jun15 2015 44.5588 -124.127 42.2

RS50_Jun15 2015 44.5602 -124.157 50.7

RS60_Jun15 2015 44.5625 -124.205 60.3

RS70_Jun15 2015 44.5588 -124.232 72.1

SB40_Jun15 2015 44.5277 -124.128 41.1

SB50_Jun15 2015 44.526 -124.165 50

SB60_Jun15 2015 44.526 -124.205 59.4

SB70_Jun15 2015 44.526 -124.235 70.5

SC001_Aug10 2010 43.607 -124.309 100

SC005_Aug10 2010 43.6037 -124.448 133

SC009_Aug10 2010 43.599 -124.363 112

SC017_Oct10 2010 43.556 -124.309 91

SC021_Aug10 2010 43.5773 -124.443 130

SC029_Aug10 2010 43.6014 -124.408 124

SC033_Aug10 2010 43.5476 -124.443 126

SC037_Oct10 2010 43.5478 -124.331 97

SC041_Aug10 2010 43.5385 -124.390 113

SC065_Aug10 2010 43.6116 -124.331 106

SC069_Aug10 2010 43.592 -124.378 117

SC073_Aug10 2010 43.5854 -124.419 125

SC077_Oct10 2010 43.5533 -124.348 104

SC097_Aug10 2010 43.5477 -124.379 112

SC105_Aug10 2010 43.5857 -124.375 108

SC128_Aug10 2010 43.5741 -124.403 119

SC134_Aug10 2010 43.5758 -124.440 129

SC136_Aug10 2010 43.5828 -124.353 110

SC150_Aug10 2010 43.6123 -124.415 127

SC401_Jun11 2011 43.6228 -124.235 24.9

SC402_Jun11 2011 43.569 -124.252 28.6
47

Station Year Latitude Longitude Depth

SC403_Jun11 2011 43.5788 -124.251 31.3

SC404_Jun11 2011 43.5957 -124.244 29.8

SC405_Jun11 2011 43.5765 -124.255 35.1

SC406_Jun11 2011 43.5508 -124.268 40.9

SC407_Jun11 2011 43.6057 -124.252 33.9

SC408_Jun11 2011 43.538 -124.278 52.5

SC409_Jun11 2011 43.5465 -124.272 47.1

SC410_Jun11 2011 43.5772 -124.267 60.8

SC411_Jun11 2011 43.602 -124.260 59.2

SC412_Jun11 2011 43.5725 -124.267 55.9

SC413_Jun11 2011 43.5953 -124.255 68.6

SC414_Jun11 2011 43.5507 -124.282 69.9

SC415_Jun11 2011 43.5805 -124.269 66

SC416_Jun11 2011 43.5512 -124.287 80.5

SC417_Jun11 2011 43.5923 -124.271 74.6

SC418_Jun11 2011 43.608 -124.270 78.4

SC419_Jun11 2011 43.6058 -124.283 89.5

SC420_Jun11 2011 43.5472 -124.297 81

SC421_Jun11 2011 43.5725 -124.285 82.7

SR30_Jun15 2015 44.4915 -124.114 31.2

SR40_Jun15 2015 44.492 -124.125 38.3

SR60_Jun15 2015 44.4943 -124.210 60

SR70_Jun15 2015 44.494 -124.251 70.1

WFP102016 2016 43.43728 -124.711 466

WFP112016 2016 43.4374 -124.683 373

WFP12016 2016 43.40523 -124.696 408

WFP122016 2016 43.458 -124.698 450

WFP132016 2016 43.46337 -124.633 243

WFP142016 2016 43.43862 -124.734 525

WFP152016 2016 43.44755 -124.721 510

WFP22016 2016 43.40605 -124.71 451

WFP32016 2016 43.4132 -124.681 348

WFP42016 2016 43.41298 -124.681 315

WFP52016 2016 43.41452 -124.727 490

WFP62016 2016 43.41447 -124.701 445

WFP72016 2016 43.41602 -124.696 406

WFP82016 2016 43.42633 -124.700 421

WFP92016 2016 43.436 -124.725 501
48

Table A-2. Abundance of all Annelids collected at the 404 stations considered in Chapter 3.

Class Subclass/Order Family Taxon

Total

Abund

Clitellata Rhynchobdellida Piscicolidae Piscicolidae sp 13

Clitellata Oligochaeta Oligochaeta spp 1

Polychaeta Echiuroidea Bonellidae Bonellia sp 27

Polychaeta Echiuroidea Echiuridae Echiura spp 1

Polychaeta Echiuroidea Echiuridae Echiuridae spp 22

Polychaeta Echiuroidea Echiuridae Echiurus echiurus 4

Polychaeta Echiuroidea Echiuridae Echiurus echiurus alascanus 6

Polychaeta Terebellida Acrocirridae Macrochaeta pege 2

Polychaeta Terebellida Ampharetidae Amage anops 5

Polychaeta Terebellida Ampharetidae Ampharete acutifrons 3

Polychaeta Terebellida Ampharetidae Ampharete finmarchica 9

Polychaeta Terebellida Ampharetidae Ampharete spp 17

Polychaeta Terebellida Ampharetidae Ampharetidae spp 10

Polychaeta Terebellida Ampharetidae Amphicteis mucronata 4

Polychaeta Terebellida Ampharetidae Amphicteis spp 6

Polychaeta Terebellida Ampharetidae Amphisamytha bioculata 1

Polychaeta Terebellida Ampharetidae Anobothrus gracilis 23

Polychaeta Terebellida Ampharetidae Asabellides lineata 4

Polychaeta Terebellida Ampharetidae Lysippe labiata 7

Polychaeta Terebellida Ampharetidae Melinna heterodonta 20

Polychaeta Terebellida Ampharetidae Melinna oculata 6

Polychaeta Terebellida Ampharetidae Melinna spp 3

Polychaeta Terebellida Ampharetidae Samytha californiensis 4

Polychaeta Amphinomida Amphinomidae Amphinomidae sp 2

Polychaeta Amphinomida Amphinomidae Chloeia pinnata 150

Polychaeta Phyllodocida Aphroditidae Aphrodita refulgida 3

Polychaeta Phyllodocida Aphroditidae Aphrodita spp 1

Polychaeta Spionida Apistobranchidae Apistobranchus tullbergi 10

Polychaeta Sedentaria Capitellidae Barantolla americana 1

Polychaeta Sedentaria Capitellidae Capitellidae spp 92

Polychaeta Sedentaria Capitellidae Decamastus gracilis 7

Polychaeta Sedentaria Capitellidae Heteromastus filobranchus 2

Polychaeta Sedentaria Capitellidae Mediomastus californiensis 248

Polychaeta Sedentaria Capitellidae Notomastus hemipodus 158

Polychaeta Sedentaria Capitellidae Notomastus latericeus 152

Polychaeta Sedentaria Capitellidae Notomastus lineatus 90

Polychaeta Sedentaria Capitellidae Notomastus magnus 2

Polychaeta Sedentaria Capitellidae Notomastus spp 114

Polychaeta Sedentaria Capitellidae Notomastus tenuis 145

Polychaeta Sedentaria Chaetopteridae Chaetopteridae sp 2

Polychaeta Sedentaria Chaetopteridae Chaetopterus sp 1
49

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Sedentaria Chaetopteridae Chaetopterus variopedatus cmplx 1

Polychaeta Sedentaria Chaetopteridae Mesochaetopterus spp 3

Polychaeta Sedentaria Chaetopteridae Mesochaetopterus taylori 26

Polychaeta Sedentaria Chaetopteridae Phyllochaetopterus limicolus 3

Polychaeta Sedentaria Chaetopteridae Phyllochaetopterus prolifica 1

Polychaeta Sedentaria Chaetopteridae Spiochaetopterus costarum 618

Polychaeta Sedentaria Chaetopteridae Spiochaetopterus spp 25

Polychaeta Terebellida Cirratulidae Aphelochaeta spp 237

Polychaeta Terebellida Cirratulidae Aphelochaeta tigrina 18

Polychaeta Terebellida Cirratulidae Chaetozone bansei 124

Polychaeta Terebellida Cirratulidae Chaetozone columbiana 3

Polychaeta Terebellida Cirratulidae Chaetozone nr setosa 3

Polychaeta Terebellida Cirratulidae Chaetozone spp 19

Polychaeta Terebellida Cirratulidae Cirratulidae spp 5

Polychaeta Terebellida Cirratulidae Kirkegaardia dutchae 3

Polychaeta Terebellida Cirratulidae Kirkegaardia secunda 4

Polychaeta Terebellida Cirratulidae Kirkegaardia serratiseta 2

Polychaeta Terebellida Cirratulidae Kirkegaardia tesselata 29

Polychaeta Terebellida Cirratulidae Monticellina cryptica 5

Polychaeta Terebellida Cirratulidae Monticellina serratiseta 2

Polychaeta Terebellida Cirratulidae Monticellina spp 3

Polychaeta Terebellida Cirratulidae Monticellina tesselata 59

Polychaeta Sedentaria Cossuridae Cossura bansei 1

Polychaeta Sedentaria Cossuridae Cossura candida 11

Polychaeta Sedentaria Cossuridae Cossura pygodactylata 1

Polychaeta Eunicida Dorvilleidae Dorvillea annulata 7

Polychaeta Eunicida Dorvilleidae Protodorvillea gracilis 8

Polychaeta Eunicida Dorvilleidae Schistomeringos longicornis 3

Polychaeta Terebellida Fauveliopsidae Fauveliopsis magna 6

Polychaeta Terebellida Flabelligeridae Brada sachalina 15

Polychaeta Terebellida Flabelligeridae Brada villosa 13

Polychaeta Terebellida Flabelligeridae Flabelligera affinis 1

Polychaeta Terebellida Flabelligeridae Flabelligeridae spp 1

Polychaeta Terebellida Flabelligeridae Pherusa neopapillata 5

Polychaeta Terebellida Flabelligeridae Pherusa plumosa 3

Polychaeta Terebellida Flabelligeridae Piromis hospitis 1

Polychaeta Phyllodocida Glyceridae Glycera americana 5

Polychaeta Phyllodocida Glyceridae Glycera macrobranchia 2

Polychaeta Phyllodocida Glyceridae Glycera nana 103

Polychaeta Phyllodocida Glyceridae Glycera oxycephala 117

Polychaeta Phyllodocida Glyceridae Glycera spp 6

Polychaeta Phyllodocida Glyceridae Glycera tesselata 1
50

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Phyllodocida Goniadidae Glycinde armigera 239

Polychaeta Phyllodocida Goniadidae Glycinde picta 21

Polychaeta Phyllodocida Goniadidae Glycinde spp 6

Polychaeta Phyllodocida Goniadidae Goniada brunnea 15

Polychaeta Phyllodocida Goniadidae Goniada maculata 57

Polychaeta Phyllodocida Goniadidae Goniada spp 4

Polychaeta Phyllodocida Hesionidae Heteropodarke heteromorpha 2

Polychaeta Phyllodocida Hesionidae Ophiodromus pugettensis 4

Polychaeta Phyllodocida Hesionidae Podarkeopsis glabrus 7

Polychaeta Eunicida Lumbrineridae Eranno bicirrata 79

Polychaeta Eunicida Lumbrineridae Eranno lagunae 24

Polychaeta Eunicida Lumbrineridae Eranno sp 11

Polychaeta Eunicida Lumbrineridae Lumbrineridae spp 20

Polychaeta Eunicida Lumbrineridae Lumbrineris californiensis 9

Polychaeta Eunicida Lumbrineridae Lumbrineris cruzensis 23

Polychaeta Eunicida Lumbrineridae Lumbrineris japonica 7

Polychaeta Eunicida Lumbrineridae Lumbrineris latreilli 15

Polychaeta Eunicida Lumbrineridae Lumbrineris spp 85

Polychaeta Eunicida Lumbrineridae Ninoe gemmea 246

Polychaeta Eunicida Lumbrineridae Scoletoma luti 294

Polychaeta Eunicida Lumbrineridae Scoletoma tetraura 7

Polychaeta Eunicida Lumbrineridae Scoletoma zonata 4

Polychaeta Spionida Magelonidae Magelona berkeleyi 931

Polychaeta Spionida Magelonidae Magelona hartmanae 109

Polychaeta Spionida Magelonidae Magelona longicornis 180

Polychaeta Spionida Magelonidae Magelona sacculata 275

Polychaeta Spionida Magelonidae Magelona spp 2

Polychaeta Sedentaria Maldanidae Axiothella rubrocincta 119

Polychaeta Capitellida Maldanidae Chirimia similis 69

Polychaeta Sedentaria Maldanidae Clymenura gracilis 24

Polychaeta Sedentaria Maldanidae Clymenura spp 8

Polychaeta Sedentaria Maldanidae Euclymene zonalis 25

Polychaeta Sedentaria Maldanidae Euclymeninae spp 319

Polychaeta Sedentaria Maldanidae Maldane cristata 5

Polychaeta Sedentaria Maldanidae Maldane sarsi 247

Polychaeta Sedentaria Maldanidae Maldanidae spp 4

Polychaeta Sedentaria Maldanidae Metasychis disparidentatus 14

Polychaeta Sedentaria Maldanidae Nicomache lumbricalis 1

Polychaeta Sedentaria Maldanidae Nicomache personata 1

Polychaeta Sedentaria Maldanidae Notoproctus pacificus 11

Polychaeta Sedentaria Maldanidae Petaloproctus borealis 8

Polychaeta Sedentaria Maldanidae Praxillella gracilis 89
51

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Sedentaria Maldanidae Praxillella pacifica 13

Polychaeta Sedentaria Maldanidae Praxillella spp 28

Polychaeta Sedentaria Maldanidae Rhodine bitorquata 70

Polychaeta Phyllodocida Nephtyidae Bipalponephtys cornuta 9

Polychaeta Phyllodocida Nephtyidae Nephtys caeca 4

Polychaeta Phyllodocida Nephtyidae Nephtys caecoides 217

Polychaeta Phyllodocida Nephtyidae Nephtys ferruginea 146

Polychaeta Phyllodocida Nephtyidae Nephtys glabra 5

Polychaeta Phyllodocida Nephtyidae Nephtys punctata 61

Polychaeta Phyllodocida Nephtyidae Nephtys spp 233

Polychaeta Phyllodocida Nereididae Cheilonereis cyclurus 3

Polychaeta Phyllodocida Nereididae Nereididae spp 1

Polychaeta Phyllodocida Nereididae Nereis pelagica 2

Polychaeta Phyllodocida Nereididae Nereis procera 24

Polychaeta Phyllodocida Nereididae Nereis spp 4

Polychaeta Phyllodocida Nereididae Nereis zonata 33

Polychaeta Eunicida Oenonidae Drilonereis falcata 5

Polychaeta Eunicida Oenonidae Drilonereis longa 7

Polychaeta Eunicida Oenonidae Notocirrus californiensis 5

Polychaeta Eunicida Onuphidae Diopatra ornata 43

Polychaeta Eunicida Onuphidae Onuphidae spp 20

Polychaeta Eunicida Onuphidae Onuphis elegans 7

Polychaeta Eunicida Onuphidae Onuphis iridescens 647

Polychaeta Eunicida Onuphidae Onuphis spp 21

Polychaeta Eunicida Onuphidae Paradiopatra parva 98

Polychaeta Sedentaria Opheliidae Armandia brevis 1

Polychaeta Sedentaria Opheliidae Ophelia assimilis 357

Polychaeta Sedentaria Opheliidae Ophelia pulchella 16

Polychaeta Sedentaria Opheliidae Ophelina acuminata 67

Polychaeta Sedentaria Opheliidae Ophelina cylindricaudata 1

Polychaeta Sedentaria Orbiniidae Leitoscoloplos pugettensis 266

Polychaeta Sedentaria Orbiniidae Naineris grubei 2

Polychaeta Sedentaria Orbiniidae Naineris uncinata 6

Polychaeta Sedentaria Orbiniidae Phylo felix 118

Polychaeta Sedentaria Orbiniidae Scoloplos acmeceps 317

Polychaeta Sedentaria Orbiniidae Scoloplos armiger 3

Polychaeta Sabellida Oweniidae Galathowenia oculata 1347

Polychaeta Sabellida Oweniidae Myriochele gracilis 174

Polychaeta Sabellida Oweniidae Myriochele spp 1

Polychaeta Sabellida Oweniidae Owenia fusiformis 88

Polychaeta Sedentaria Paraonidae Aricidea (acmira) catherinae 4

Polychaeta Sedentaria Paraonidae Aricidea (acmira) cerrutii 4
52

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Sedentaria Paraonidae Aricidea (acmira) simplex 36

Polychaeta Sedentaria Paraonidae Aricidea (aedicira) pacifica 25

Polychaeta Sedentaria Paraonidae Aricidea (allia) antennata 8

Polychaeta Sedentaria Paraonidae Aricidea (allia) ramosa 11

Polychaeta Sedentaria Paraonidae Aricidea spp 34

Polychaeta Sedentaria Paraonidae Cirrophorus branchiatus 3

Polychaeta Sedentaria Paraonidae Levinsenia gracilis 5

Polychaeta Sedentaria Paraonidae Paradoneis spinifera 1

Polychaeta Terebellida Pectinariidae Pectinaria californiensis 199

Polychaeta Terebellida Pectinariidae Pectinaria granulata 13

Polychaeta Terebellida Pectinariidae Pectinaria spp 15

Polychaeta Phyllodocida Pholoidae Pholoe spp 39

Polychaeta Phyllodocida Phyllodocidae Eteone leptotes 1

Polychaeta Phyllodocida Phyllodocidae Eteone spp 30

Polychaeta Phyllodocida Phyllodocidae Eulalia californiensis 2

Polychaeta Phyllodocida Phyllodocidae Eumida longicornuta 2

Polychaeta Phyllodocida Phyllodocidae Hesionura coineaui difficilis 1

Polychaeta Phyllodocida Phyllodocidae Paranaitis polynoides 15

Polychaeta Phyllodocida Phyllodocidae Phyllodoce cuspidata 20

Polychaeta Phyllodocida Phyllodocidae Phyllodoce groenlandica 12

Polychaeta Phyllodocida Phyllodocidae Phyllodoce hartmanae 41

Polychaeta Phyllodocida Phyllodocidae Phyllodoce longipes 1

Polychaeta Phyllodocida Phyllodocidae Phyllodoce spp 7

Polychaeta Phyllodocida Pilargidae Ancistrosyllis groenlandica 1

Polychaeta Phyllodocida Pilargidae Hermundura fauveli 48

Polychaeta Phyllodocida Pilargidae Hermundura ocularis 9

Polychaeta Phyllodocida Pilargidae Pilargis berkeleyae 5

Polychaeta Phyllodocida Pilargidae Pilargis maculata 21

Polychaeta Phyllodocida Pilargidae Sigambra bassi 2

Polychaeta Phyllodocida Pilargidae Sigambra spp 1

Polychaeta Polychaeta incertae sedis Polygordiidae Polygordius spp 9

Polychaeta Phyllodocida Polynoidae Arcteobia anticostiensis 2

Polychaeta Phyllodocida Polynoidae Bylgides macrolepidus 7

Polychaeta Phyllodocida Polynoidae Eucranta anoculata 9

Polychaeta Phyllodocida Polynoidae Gattyana ciliata 1

Polychaeta Phyllodocida Polynoidae Harmothoe extenuata 3

Polychaeta Phyllodocida Polynoidae Harmothoe spp 3

Polychaeta Phyllodocida Polynoidae Hesperonoe complanata 9

Polychaeta Phyllodocida Polynoidae Hesperonoe laevis 6

Polychaeta Phyllodocida Polynoidae Lepidasthenia berkeleyae 13

Polychaeta Phyllodocida Polynoidae Lepidasthenia longicirrata 10

Polychaeta Phyllodocida Polynoidae Lepidonotus spiculus 2
53

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Phyllodocida Polynoidae Lepidonotus squamatus 5

Polychaeta Phyllodocida Polynoidae Malmgreniella bansei 21

Polychaeta Phyllodocida Polynoidae Malmgreniella baschi 2

Polychaeta Phyllodocida Polynoidae Malmgreniella berkeleyorum 1

Polychaeta Phyllodocida Polynoidae Malmgreniella liei 1

Polychaeta Phyllodocida Polynoidae Malmgreniella macginitiei 1

Polychaeta Phyllodocida Polynoidae Malmgreniella sanpedroensis 46

Polychaeta Phyllodocida Polynoidae Malmgreniella scriptoria 51

Polychaeta Phyllodocida Polynoidae Malmgreniella spp 7

Polychaeta Phyllodocida Polynoidae Nemidia microlepida 6

Polychaeta Phyllodocida Polynoidae Polynoidae spp 9

Polychaeta Phyllodocida Polynoidae Tenonia priops 15

Polychaeta Sabellida Sabellidae Bispira spp 1

Polychaeta Sabellida Sabellidae Chone magna 1

Polychaeta Sabellida Sabellidae Chone spp 5

Polychaeta Sabellida Sabellidae Chone veleronis 1

Polychaeta Sabellida Sabellidae Dialychone albocincta 8

Polychaeta Sabellida Sabellidae Dialychone veleronis 6

Polychaeta Sabellida Sabellidae Euchone analis 8

Polychaeta Sabellida Sabellidae Euchone spp 2

Polychaeta Sabellida Sabellidae Myxicola infundibulum 1

Polychaeta Sabellida Sabellidae Paradialychone ecaudata 14

Polychaeta Sabellida Sabellidae Parasabella rugosa 1

Polychaeta Sabellida Sabellidae Pseudopotamilla intermedia 1

Polychaeta Sabellida Sabellidae Sabellidae spp 3

Polychaeta Polychaeta incertae sedis Saccocirridae Saccocirrus spp 2

Polychaeta Sedentaria Scalibregmatidae Scalibregma californicum 77

Polychaeta Sedentaria Scalibregmatidae Scalibregma inflatum 8

Polychaeta Phyllodocida Sigalionidae Leanira alba 3

Polychaeta Phyllodocida Sigalionidae Pholoides asperus 40

Polychaeta Phyllodocida Sigalionidae Pisione spp 3

Polychaeta Phyllodocida Sigalionidae Sigalion spinosus 75

Polychaeta Phyllodocida Sigalionidae Sthenelais berkeleyi 7

Polychaeta Phyllodocida Sigalionidae Sthenelais verruculosa 12

Polychaeta Spionida Spionidae Aonides glandulosa 2

Polychaeta Spionida Spionidae Apoprionospio pygmaea 1

Polychaeta Spionida Spionidae Boccardia pugettensis 21

Polychaeta Spionida Spionidae Boccardia sp 9

Polychaeta Spionida Spionidae Dipolydora brachycephala 2

Polychaeta Spionida Spionidae Dipolydora cardalia 5

Polychaeta Spionida Spionidae Dipolydora caulleryi 2

Polychaeta Spionida Spionidae Dipolydora socialis 13
54

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Spionida Spionidae Laonice cirrata 29

Polychaeta Spionida Spionidae Microspio pigmentata 68

Polychaeta Spionida Spionidae Paraprionospio alata 246

Polychaeta Spionida Spionidae Prionospio ehlersi 8

Polychaeta Spionida Spionidae Prionospio lighti 16

Polychaeta Spionida Spionidae Prionospio steenstrupi 209

Polychaeta Spionida Spionidae Pseudopolydora sp 1

Polychaeta Spionida Spionidae Scolelepis squamata 48

Polychaeta Spionida Spionidae Scolelepis tridentata 2

Polychaeta Spionida Spionidae Spio spp 1

Polychaeta Spionida Spionidae Spio thulini 38

Polychaeta Spionida Spionidae Spionidae spp 5

Polychaeta Spionida Spionidae Spiophanes berkeleyorum 185

Polychaeta Spionida Spionidae Spiophanes norrisi 300

Polychaeta Terebellida Sternaspidae Sternaspis assimilis 65

Polychaeta Terebellida Sternaspidae Sternaspis fossor 324

Polychaeta Phyllodocida Syllidae Exogone lourei 28

Polychaeta Phyllodocida Syllidae Exogone spp 33

Polychaeta Phyllodocida Syllidae Geminosyllis ohma 2

Polychaeta Phyllodocida Syllidae Pionosyllis sp 1

Polychaeta Phyllodocida Syllidae Proceraea cornuta 1

Polychaeta Phyllodocida Syllidae Sphaerosyllis californiensis 42

Polychaeta Phyllodocida Syllidae Syllides longocirratus 3

Polychaeta Phyllodocida Syllidae Syllis spp 2

Polychaeta Phyllodocida Syllidae Typosyllis cornuta 12

Polychaeta Phyllodocida Syllidae Typosyllis heterochaeta 24

Polychaeta Phyllodocida Syllidae Typosyllis hyperioni 2

Polychaeta Phyllodocida Syllidae Typosyllis pigmentata 9

Polychaeta Phyllodocida Syllidae Typosyllis spp 10

Polychaeta Terebellida Terebellidae Amaeana occidentalis 45

Polychaeta Terebellida Terebellidae Artacama coniferi 27

Polychaeta Terebellida Terebellidae Lanassa sp 1

Polychaeta Terebellida Terebellidae Lanassa venusta 3

Polychaeta Terebellida Terebellidae Pista agassizi 3

Polychaeta Terebellida Terebellidae Pista brevibranchiata 24

Polychaeta Terebellida Terebellidae Pista estevanica 142

Polychaeta Terebellida Terebellidae Pista moorei 6

Polychaeta Terebellida Terebellidae Pista spp 27

Polychaeta Terebellida Terebellidae Pista wui 107

Polychaeta Terebellida Terebellidae Polycirrus californicus 5

Polychaeta Terebellida Terebellidae Polycirrus spp 145

Polychaeta Terebellida Terebellidae Proclea graffi 9
55

Class Subclass/Order Family Taxon

Total

Abund

Polychaeta Terebellida Terebellidae Proclea graffii 3

Polychaeta Terebellida Terebellidae Scionella japonica 2

Polychaeta Terebellida Terebellidae Streblosoma bairdi 39

Polychaeta Terebellida Terebellidae Thelepus setosus 3

Polychaeta Echiuroidea Thalassematidae Arhynchite pugettensis 16

Polychaeta Sedentaria Travisiidae Travisia brevis 41

Polychaeta Sedentaria Travisiidae Travisia forbesii 20

Polychaeta Sedentaria Travisiidae Travisia japonica 8

Polychaeta Sedentaria Travisiidae Travisia pupa 6

Polychaeta Terebellida Trichobranchidae Artacamella hancocki 9

Polychaeta Terebellida Trichobranchidae Terebellidae spp 2

Polychaeta Terebellida Trichobranchidae Terebellides californica 16

Polychaeta Terebellida Trichobranchidae Terebellides reishi 29

Polychaeta Terebellida Trichobranchidae Terebellides spp 21

Polychaeta Terebellida Trichobranchidae Trichobranchus glacialis 3

Polychaeta Spionida Trochochaetidae Trochochaeta multisetosa 4

Polychaeta Polychaete spp 1
56

Table A-3. Abundance of all Crustaceans collected at the 404 stations considered in Chapter 3.

Class Subclass/Order Family Taxon Total 

Abund

Hexanauplia Calanoida Calanidae Calanus spp 51

Hexanauplia Calanoida Metridinidae Metridia pacifica 1

Hexanauplia Calanoida Pontellidae Epilabidocera longipedata 7

Hexanauplia Cyclopoida Clausidiidae Hemicyclops sp 27

Hexanauplia Cyclopoida Poecilostomatoida sp 1

Hexanauplia Sessilia Balanidae Balanus crenatus 1

Hexanauplia Sessilia Balanidae Balanus sp 1

Malacostraca Amphipoda Acidostomatidae Acidostoma sp 2

Malacostraca Amphipoda Ampeliscidae Ampelisca agassizi 41

Malacostraca Amphipoda Ampeliscidae Ampelisca brevisimulata 5

Malacostraca Amphipoda Ampeliscidae Ampelisca careyi 364

Malacostraca Amphipoda Ampeliscidae Ampelisca cristata 29

Malacostraca Amphipoda Ampeliscidae Ampelisca hancocki 13

Malacostraca Amphipoda Ampeliscidae Ampelisca lobata 2

Malacostraca Amphipoda Ampeliscidae Ampelisca pugetica 7

Malacostraca Amphipoda Ampeliscidae Ampelisca spp 6

Malacostraca Amphipoda Ampeliscidae Ampelisca unsocalae 10

Malacostraca Amphipoda Ampeliscidae Ampeliscidae sp 1

Malacostraca Amphipoda Ampeliscidae Byblis millsi 1

Malacostraca Amphipoda Ampeliscidae Byblis mulleni 2

Malacostraca Amphipoda Ampeliscidae Byblis spp 4

Malacostraca Amphipoda Aoridae Aoroides inermis 20

Malacostraca Amphipoda Aoridae Aoroides sp 10

Malacostraca Amphipoda Aoridae Aoroides spp 4

Malacostraca Amphipoda Argissidae Argissa hamatipes 7

Malacostraca Amphipoda Caprellidae Caprella mendax 13

Malacostraca Amphipoda Caprellidae Caprella spp 1

Malacostraca Amphipoda Caprellidae Tritella pilimana 1

Malacostraca Amphipoda Caprellidae Tritella tenuissima 17

Malacostraca Amphipoda Corophiidae Cheirimedeia macrocarpa 13

Malacostraca Amphipoda Corophiidae Cheirimedeia macrodactyla 16

Malacostraca Amphipoda Corophiidae Cheirimedeia spp 2

Malacostraca Amphipoda Corophiidae Cheirimedeia zotea 26

Malacostraca Amphipoda Corophiidae Laticorophium baconi 4

Malacostraca Amphipoda Corophiidae Protomedeia articulata 13

Malacostraca Amphipoda Corophiidae Protomedeia prudens 28

Malacostraca Amphipoda Corophiidae Protomedeia spp 2

Malacostraca Amphipoda Dulichiidae Dulichia sp 1

Malacostraca Amphipoda Eusiridae Rhachotropis sp 1

Malacostraca Amphipoda Eusiridae Rhacotropis barnardi 1

Malacostraca Amphipoda Eusiridae Rhacotropis boreopacifica 1

Malacostraca Amphipoda Haustoriidae Eohaustorius sawyeri 181

Malacostraca Amphipoda Haustoriidae Eohaustorius sencillus 158

Malacostraca Amphipoda Hyperiidae Hyperiidae sp 5

Malacostraca Amphipoda Isaeidae Isaeidae spp 79

Malacostraca Amphipoda Ischyroceridae Ischyrocerus spp 2
57

Class Subclass/Order Family Taxon Total 

Abund

Malacostraca Amphipoda Liljeborgiidae Idunella sp 4

Malacostraca Amphipoda Liljeborgiidae Liljeborgia sp 10

Malacostraca Amphipoda Liljeborgiidae Listriella albina 3

Malacostraca Amphipoda Lysianassidae Hippomedon coecus 2

Malacostraca Amphipoda Lysianassidae Hippomedon columbianus 1

Malacostraca Amphipoda Lysianassidae Hippomedon spp 1

Malacostraca Amphipoda Lysianassidae Lepidepecreum garthi 6

Malacostraca Amphipoda Lysianassidae Lysianassidae spp 11

Malacostraca Amphipoda Lysianassidae Orchomenella decipiens 1

Malacostraca Amphipoda Lysianassidae Orchomenella pacifica 18

Malacostraca Amphipoda Lysianassidae Orchomenella pinguis 115

Malacostraca Amphipoda Lysianassidae Psammonyx longimerus 4

Malacostraca Amphipoda Lysianassidae Wecomedon wecomus 9

Malacostraca Amphipoda Maeridae Maera loveni 1

Malacostraca Amphipoda Maeridae Maera sp 1

Malacostraca Amphipoda Megaluropidae Gibberosus myersi 42

Malacostraca Amphipoda Melitidae Desdimelita desdichada 6

Malacostraca Amphipoda Melitidae Melitidae sp 5

Malacostraca Amphipoda Oedicerotidae Americhelidium pectinatum 1

Malacostraca Amphipoda Oedicerotidae Americhelidium shoemakeri 15

Malacostraca Amphipoda Oedicerotidae Bathymedon covilhani 1

Malacostraca Amphipoda Oedicerotidae Bathymedon pumilis 1

Malacostraca Amphipoda Oedicerotidae Hartmanodes hartmanae 1

Malacostraca Amphipoda Oedicerotidae Monoculodes emarginatus 3

Malacostraca Amphipoda Oedicerotidae Monoculodes latissimus 1

Malacostraca Amphipoda Oedicerotidae Oedicerotidae spp 2

Malacostraca Amphipoda Oedicerotidae Pacifoculodes spinipes 36

Malacostraca Amphipoda Oedicerotidae Westwoodilla tone 18

Malacostraca Amphipoda Opisidae Opisa tridentata 2

Malacostraca Amphipoda Pachynidae Prachynella lodo 4

Malacostraca Amphipoda Pardaliscidae Nicippe tumida 7

Malacostraca Amphipoda Pardaliscidae Pardaliscella symmetrica 2

Malacostraca Amphipoda Photidae Gammaropsis ellisi 1

Malacostraca Amphipoda Photidae Gammaropsis spp 7

Malacostraca Amphipoda Photidae Photis brevipes 81

Malacostraca Amphipoda Photidae Photis chiconola 1

Malacostraca Amphipoda Photidae Photis macinerneyi 14

Malacostraca Amphipoda Photidae Photis spp 21

Malacostraca Amphipoda Phoxocephalidae Foxiphalus golfensis 2

Malacostraca Amphipoda Phoxocephalidae Foxiphalus similis 12

Malacostraca Amphipoda Phoxocephalidae Foxiphalus xiximeus 4

Malacostraca Amphipoda Phoxocephalidae Grandifoxus longirostris 24

Malacostraca Amphipoda Phoxocephalidae Harpiniopsis emeryi 7

Malacostraca Amphipoda Phoxocephalidae Harpiniopsis fulgens 5

Malacostraca Amphipoda Phoxocephalidae Harpiniopsis spp 8

Malacostraca Amphipoda Phoxocephalidae Heterophoxus affinis 60

Malacostraca Amphipoda Phoxocephalidae Heterophoxus ellisi 11
58

Class Subclass/Order Family Taxon Total 

Abund

Malacostraca Amphipoda Phoxocephalidae Heterophoxus frequens 1

Malacostraca Amphipoda Phoxocephalidae Heterophoxus oculatus 1

Malacostraca Amphipoda Phoxocephalidae Heterophoxus spp 121

Malacostraca Amphipoda Phoxocephalidae Majoxiphalus major 152

Malacostraca Amphipoda Phoxocephalidae Mandibulophoxus mayi 45

Malacostraca Amphipoda Phoxocephalidae Metaphoxus frequens 9

Malacostraca Amphipoda Phoxocephalidae Paraphoxus sp 2

Malacostraca Amphipoda Phoxocephalidae Pseudharpina inexpectata 1

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius abronius 10

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius bicuspidatus 28

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius boreovariatus 67

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius dabouis 63

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius fatigans 214

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius spp 317

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius variatus 56

Malacostraca Amphipoda Phoxocephalidae Rhepoxynius vigitegus 276

Malacostraca Amphipoda Pleustidae Kamptopleustes coquillus 2

Malacostraca Amphipoda Pleustidae Pleusymtes spp 4

Malacostraca Amphipoda Stenothoidae Stenothoidae spp 6

Malacostraca Amphipoda Synopiidae Bruzelia tuberculata 1

Malacostraca Amphipoda Synopiidae Syrrhoe longifrons 5

Malacostraca Amphipoda Uristidae Anonyx sp 1

Malacostraca Cumacea Diastylidae Anchicolurus occidentalis 33

Malacostraca Cumacea Diastylidae Diastylidae spp 2

Malacostraca Cumacea Diastylidae Diastylis abboti 1

Malacostraca Cumacea Diastylidae Diastylis bidentata 3

Malacostraca Cumacea Diastylidae Diastylis dalli 1

Malacostraca Cumacea Diastylidae Diastylis paraspinulosa 7

Malacostraca Cumacea Diastylidae Diastylis pellucida 9

Malacostraca Cumacea Diastylidae Diastylis quadriplicata 3

Malacostraca Cumacea Diastylidae Diastylis santamariensis 13

Malacostraca Cumacea Diastylidae Diastylis spp 5

Malacostraca Cumacea Diastylidae Diastylopsis dawsoni 33

Malacostraca Cumacea Diastylidae Diastylopsis tenuis 3

Malacostraca Cumacea Lampropidae Hemilamprops californicus 14

Malacostraca Cumacea Leuconidae Eudorella emarginata 7

Malacostraca Cumacea Leuconidae Eudorella pacifica 26

Malacostraca Cumacea Leuconidae Eudorellopsis longirostris 7

Malacostraca Cumacea Leuconidae Leucon spp 3

Malacostraca Cumacea Leuconidae Leuconidae sp 3

Malacostraca Cumacea Nannastacidae Campylaspis hartae 1

Malacostraca Cumacea Nannastacidae Campylaspis papillata 3

Malacostraca Cumacea Nannastacidae Campylaspis rubromaculata 1

Malacostraca Cumacea Nannastacidae Campylaspis rufa 1

Malacostraca Cumacea Nannastacidae Campylaspis spp 6

Malacostraca Cumacea Cumacea spp 1

Malacostraca Decapoda Axiidae Calocarides (Acanthaxius) 

spinulicauda

2
59

Class Subclass/Order Family Taxon Total 

Abund

Malacostraca Decapoda Callianassidae Neotrypaea californiensis 18

Malacostraca Decapoda Callianassidae Neotrypaea gigas 10

Malacostraca Decapoda Callianassidae Neotrypaea spp 99

Malacostraca Decapoda Cancridae Metacarcinus spp 8

Malacostraca Decapoda Cancridae Romaleon jordani 1

Malacostraca Decapoda Caridea Caridea spp 15

Malacostraca Decapoda Crangonidae Crangon alaskensis 13

Malacostraca Decapoda Crangonidae Crangon franciscorum 1

Malacostraca Decapoda Crangonidae Crangon spp 21

Malacostraca Decapoda Crangonidae Crangon stylirostris 16

Malacostraca Decapoda Crangonidae Lissocrangon stylirostris 14

Malacostraca Decapoda Crangonidae Neocrangon communis 6

Malacostraca Decapoda Diogenidae Paguristes turgidus 1

Malacostraca Decapoda Oregoniidae Oregonia gracilis 1

Malacostraca Decapoda Paguridae Pagurus armatus 15

Malacostraca Decapoda Paguridae Pagurus capillatus 1

Malacostraca Decapoda Paguridae Pagurus caurinus 1

Malacostraca Decapoda Paguridae Pagurus dalli 1

Malacostraca Decapoda Paguridae Pagurus ochotensis 4

Malacostraca Decapoda Paguridae Pagurus quaylei 6

Malacostraca Decapoda Paguridae Pagurus spp 33

Malacostraca Decapoda Pandalidae Pandalus jordani 3

Malacostraca Decapoda Pasiphaeidae Pasiphaea pacifica 1

Malacostraca Decapoda Pinnotheridae Pinnixa occidentalis complex 1041

Malacostraca Decapoda Pinnotheridae Pinnixa spp 162

Malacostraca Decapoda Pinnotheridae Pinnotheridae sp 17

Malacostraca Decapoda Thoridae Eualus berkeleyorum 1

Malacostraca Decapoda Thoridae Spirontocaris lamellicornis 1

Malacostraca Decapoda Thoridae Spirontocaris snyderi 2

Malacostraca Decapoda Thoridae Spirontocaris sp 2

Malacostraca Euphausiacea Euphausiidae Euphausia pacifica 2

Malacostraca Euphausiacea Euphausiidae Thysanoessa spp 2

Malacostraca Isopoda Aegidae Rocinela angustata 2

Malacostraca Isopoda Ancinidae Bathycopea daltonae 8

Malacostraca Isopoda Anthuridae Haliophasma geminata 2

Malacostraca Isopoda Gnathiidae Gnathia spp 1

Malacostraca Isopoda Gnathiidae Gnathia tridens 1

Malacostraca Isopoda Idoteidae Edotia sublittoralis 4

Malacostraca Isopoda Idoteidae Synidotea media 3

Malacostraca Isopoda Idoteidae Synidotea spp 21

Malacostraca Leptostraca Nebaliidae Nebalia spp 7

Malacostraca Mysida Mysidae Alienacanthomysis macropsis 2

Malacostraca Mysida Mysidae Archaeomysis grebnitzkii 96

Malacostraca Mysida Mysidae Exacanthomysis davisi 1

Malacostraca Mysida Mysidae Holmesiella anomala 2

Malacostraca Mysida Mysidae Inusitatomysis insolita 1

Malacostraca Mysida Mysidae Mysidae sp 6
60

Class Subclass/Order Family Taxon Total 

Abund

Malacostraca Mysida Mysidae Mysidella americana 1

Malacostraca Mysida Mysidae Neomysis kadiakensis 2

Malacostraca Mysida Mysidae Pacifacanthomysis 

nephrophthalma

3

Malacostraca Tanaidacea Paratanaoidea incertae sedis Scoloura phillipsi 3

Malacostraca Tanaidacea Tanaellidae Araphura breviaria 1

Ostracoda Myodocopida Cylindroleberididae Cylindroleberididae spp 43

Ostracoda Myodocopida Philomedidae Euphilomedes carcharadonta 22

Ostracoda Myodocopida Philomedidae Euphilomedes carcharodonta 262

Ostracoda Myodocopida Philomedidae Euphilomedes producta 29

Ostracoda Myodocopida Philomedidae Scleroconcha trituberculata 22

Ostracoda Myodocopida Rutidermatidae Rutiderma lomae 3

Ostracoda Ostracoda spp 4
61

Table A-4. Abundance of all Molluscs collected at the 404 stations considered in Chapter 3.

Class Subclass/Order Family Taxon Total

Abund

Bivalvia Adapedonta Hiatellidae Hiatella arctica 14

Bivalvia Adapedonta Pharidae Pharidae sp 2

Bivalvia Adapedonta Pharidae Siliqua alta 7

Bivalvia Adapedonta Pharidae Siliqua patula 24

Bivalvia Adapedonta Pharidae Siliqua spp 4

Bivalvia Adapedonta Solenidae Solen sicarius 1

Bivalvia Cardiida Cardiidae Keenaea centifilosa 6

Bivalvia Cardiida Tellinidae Ameritella carpenteri 143

Bivalvia Cardiida Tellinidae Ameritella modesta 60

Bivalvia Cardiida Tellinidae Macoma calcarea 1

Bivalvia Cardiida Tellinidae Macoma carlottensis 725

Bivalvia Cardiida Tellinidae Macoma elimata 223

Bivalvia Cardiida Tellinidae Macoma inquinata 1

Bivalvia Cardiida Tellinidae Macoma spp 62

Bivalvia Cardiida Tellinidae Macoma yoldiformis 1

Bivalvia Cardiida Tellinidae Megangulus bodegensis 17

Bivalvia Cardiida Tellinidae Tellina nuculoides 230

Bivalvia Carditida Carditidae Cyclocardia ventricosa 140

Bivalvia Galeommatida Lasaeidae Kurtiella tumida 868

Bivalvia Heterodonta Cuspidariidae Cardiomya pectinata 13

Bivalvia Heterodonta Cuspidariidae Cardiomya planetica 3

Bivalvia Heterodonta Cuspidariidae Cuspidaria glacialis 4

Bivalvia Heterodonta Cuspidariidae Cuspidariidae sp 3

Bivalvia Heterodonta Cuspidariidae Pseudoneaera (Austroneaera) 

semipellucida

1

Bivalvia Heterodonta Laternulidae Laternula marilina 28

Bivalvia Heterodonta Lyonsiidae Lyonsia californica 3

Bivalvia Heterodonta Pandoridae Pandora bilirata 20

Bivalvia Heterodonta Pandoridae Pandora filosa 2

Bivalvia Heterodonta Thraciidae Thracia challisiana 2

Bivalvia Heterodonta Thraciidae Thracia trapezoides 6

Bivalvia Heterodonta Basterotiidae Saxicavella pacifica 25

Bivalvia Lucinida Lucinidae Lucinoma annulata 15

Bivalvia Lucinida Lucinidae Lucinoma annulatum 2

Bivalvia Lucinida Lucinidae Parvilucina tenuisculpta 1

Bivalvia Lucinida Thyasiridae Adontorhina cyclia 394

Bivalvia Lucinida Thyasiridae Adontorhina sphaericosa 31

Bivalvia Lucinida Thyasiridae Axinopsida serricata 8806

Bivalvia Lucinida Thyasiridae Mendicula ferruginosa 41

Bivalvia Lucinida Thyasiridae Thyasira flexuosa 110

Bivalvia Myida Pholadoidea Pholadoidea sp 12

Bivalvia Mytilida Mytilidae Modiolus rectus 5
62

Class Subclass/Order Family Taxon Total

Abund

Bivalvia Mytilida Mytilidae Mytilidae spp 4

Bivalvia Mytilida Mytilidae Solamen columbianum 60

Bivalvia Nuculanida Nuculanidae Nuculana conceptionis 2

Bivalvia Nuculanida Nuculanidae Nuculana fossa 17

Bivalvia Nuculanida Nuculanidae Nuculana hamata 23

Bivalvia Nuculanida Nuculanidae Nuculana minuta 1

Bivalvia Nuculanida Nuculanidae Nuculana spp 3

Bivalvia Nuculanida Yoldiidae Megayoldia martyria 22

Bivalvia Nuculanida Yoldiidae Megayoldia thraciaeformis 2

Bivalvia Nuculanida Yoldiidae Yoldia scissurata 15

Bivalvia Nuculanida Yoldiidae Yoldia seminuda 51

Bivalvia Nuculanida Yoldiidae Yoldiella spp 3

Bivalvia Nuculanida Yoldiidae Yolida scissurata 14

Bivalvia Nuculida Nuculidae Acila castrensis 1311

Bivalvia Nuculida Nuculidae Ennucula tenuis 903

Bivalvia Pectinida Pectinidae Delectopecten vancouverensis 1

Bivalvia Solemyida Nucinellidae Huxleyia munita 1171

Bivalvia Solemyida Solemyidae Solemya reidi 11

Bivalvia Venerida Mactridae Mactridae spp 1

Bivalvia Venerida Veneridae Compsomyax subdiaphana 85

Bivalvia Venerida Veneridae Humilaria kennerleyi 2

Bivalvia Venerida Veneridae Nutricola lordi 4041

Bivalvia Bivalvia spp 11

Caudofoveata Chaetodermatida Chaetodermatidae Chaetoderma argenteum 135

Caudofoveata Chaetodermatida Chaetodermatidae Chaetoderma spp 28

Gastropoda Caenogastropoda Cerithiidae Lirobittium sp 38

Gastropoda Caenogastropoda Epitoniidae Epitonium hindsii 1

Gastropoda Caenogastropoda Epitoniidae Epitonium indianorum 7

Gastropoda Caenogastropoda Epitoniidae Epitonium spp 15

Gastropoda Cephalaspidea Acteocinidae Acteocina cerealis 6

Gastropoda Cephalaspidea Acteocinidae Acteocina eximia 121

Gastropoda Cephalaspidea Acteocinidae Acteocina spp 160

Gastropoda Cephalaspidea Aglajidae Melanochlamys diomedea 6

Gastropoda Cephalaspidea Cylichnidae Cylichna attonsa 547

Gastropoda Cephalaspidea Cylichnidae Cylichna spp 1

Gastropoda Cephalaspidea Diaphanidae Diaphana californica 8

Gastropoda Cephalaspidea Gastropteridae Gastropteron pacificum 9

Gastropoda Cephalaspidea Haminoeidae Haminoea vesicula 7

Gastropoda Cephalaspidea Retusidae Retusa xystrum 1

Gastropoda Cephalaspidea Rhizoridae Volvulella cylindrica 23

Gastropoda Heterobranchia Acteonidae Rictaxis punctocaelatus 17

Gastropoda Heterobranchia Pyramidellidae Boonea oregensis 18
63

Class Subclass/Order Family Taxon Total

Abund

Gastropoda Heterobranchia Pyramidellidae Odostomia spp 187

Gastropoda Heterobranchia Pyramidellidae Turbonilla spp 116

Gastropoda Lepetellida Fissurellidae Fissurellidae sp 1

Gastropoda Lepetellida Scissurellidae Anatoma (Scissurella) crispata 1

Gastropoda Littorinimorpha Eulimidae Polygireulima rutila 9

Gastropoda Littorinimorpha Littorinidae Lacuna vincta 2

Gastropoda Littorinimorpha Naticidae Cryptonatica affinis 9

Gastropoda Littorinimorpha Naticidae Sinum scopulosum 1

Gastropoda Littorinimorpha Rissoidae Alvania compacta 50

Gastropoda Neogastropoda Borsoniidae Ophiodermella cancellata 4

Gastropoda Neogastropoda Buccinidae Ancistrolepis eucosmius 4

Gastropoda Neogastropoda Buccinidae Buccinidae sp 2

Gastropoda Neogastropoda Buccinidae Buccinum strigillatum fucanum 7

Gastropoda Neogastropoda Buccinidae Colus halli 3

Gastropoda Neogastropoda Cancellariidae Admete gracilior 2

Gastropoda Neogastropoda Cancellariidae Admete viridula 16

Gastropoda Neogastropoda Columbellidae Mitrella (Alia/Astyris) aurantiaca 35

Gastropoda Neogastropoda Columbellidae Mitrella (Alia/Astyris) gausapata 729

Gastropoda Neogastropoda Mangeliidae Kurtzia arteaga 47

Gastropoda Neogastropoda Mangeliidae Kurtziella plumbea 10

Gastropoda Neogastropoda Mangeliidae Oenopota fidicula 14

Gastropoda Neogastropoda Mangeliidae Oenopota spp 3

Gastropoda Neogastropoda Mangeliidae Oenopota turricula 1

Gastropoda Neogastropoda Muricidae Boreotrophon dalli 1

Gastropoda Neogastropoda Muricidae Boreotrophon orpheus 5

Gastropoda Neogastropoda Muricidae Scabrotrophon lasius 2

Gastropoda Neogastropoda Nassariidae Hima mendica 1

Gastropoda Neogastropoda Nassariidae Nassarius fossatus (Caesia fossata) 17

Gastropoda Neogastropoda Olividae Callianax baetica 363

Gastropoda Neogastropoda Olividae Callianax biplicata 37

Gastropoda Neogastropoda Olividae Callianax pycna 293

Gastropoda Neogastropoda Olividae Olivella baetica 24

Gastropoda Neogastropoda Pseudomelatomidae Antiplanes catalinae (voyi) 5

Gastropoda Neogastropoda Pseudomelatomidae Antiplanes santarosana 1

Gastropoda Neogastropoda Pseudomelatomidae Antiplanes spp 3

Gastropoda Neogastropoda Ptychatractidae Exilioidea rectirostris 1

Gastropoda Neogastropoda Turridae Turridae spp 5

Gastropoda Nudibranchia Arminidae Armina californica 1

Gastropoda Patellogastropoda Lottiidae Lottia sp 2

Gastropoda Trochida Colloniidae Homalopoma radiatum 9

Gastropoda Trochida Margaritidae Margarites pupillus 1

Gastropoda Trochida Solariellidae Solariella peramabilis 13
64

Class Subclass/Order Family Taxon Total

Abund

Gastropoda Trochida Solariellidae Solariella sp 1

Gastropoda Trochida Trochidae Lirularia lirulata 8

Gastropoda Trochida Trochidae Trochidae sp 3

Gastropoda Gastropoda spp 5

Polyplacophora Chitonida Ischnochitonidae Ischnochiton willetti 21

Polyplacophora Chitonida Ischnochitonidae Tripoplax trifida 5

Polyplacophora Chitonida Mopaliidae Mopalia imporcata 1

Polyplacophora Chitonida Mopaliidae Mopalia sp 2

Polyplacophora Polyplacophora sp 1

Scaphopoda Dentaliida Dentaliidae Antalis pretiosum 45

Scaphopoda Dentaliida Dentaliidae Dentaliidae spp 4

Scaphopoda Dentaliida Rhabdidae Rhabdus rectius 562

Scaphopoda Gadilida Gadilidae Gadila aberrans 105

Scaphopoda Gadilida Gadilidae Gadila tolmiei 164

Scaphopoda Gadilida Pulsellidae Pulsellidae sp 84

Scaphopoda Gadilida Pulsellidae Pulsellum salishorum 266

Scaphopoda Scaphopoda spp 4
65

Table A-5. Abundance of all other organisms not listed above collected at the 404 stations 

considered in Chapter 3.

Phylum Class Subclass/Order Family Taxon Total 

Abund

Brachiopoda Rhynchonellata Terebratulida Laqueidae Laqueus californianus 4

Bryozoa Gymnolaemata Ctenostomatida Clavoporidae Clavopora 

occidentalis

1

Chaetognatha Sagittoidea Aphragmophora Sagittidae Sagitta spp 4

Chordata Ascidacea Phlebobranchia Corellidae Chelysoma 

columbianum

12

Cnidaria Anthozoa Actiniaria Edwardsiidae Edwardsia juliae 3

Cnidaria Anthozoa Actiniaria Edwardsiidae Edwardsia profunda 1

Cnidaria Anthozoa Actiniaria Edwardsiidae Edwardsia spp 12

Cnidaria Anthozoa Actiniaria Halcampidae Halcampa crypta 2

Cnidaria Anthozoa Actiniaria Halcampidae Halcampa 

decemtentaculata

19

Cnidaria Anthozoa Actiniaria Halcampidae Halcampidae sp 9

Cnidaria Anthozoa Actiniaria Halcampidae Halcampoides

purpureus

2

Cnidaria Anthozoa Actiniaria Haloclavidae Peachia 

quinquecapitata

11

Cnidaria Anthozoa Actiniaria Metridiidae Metridium sp 12

Cnidaria Anthozoa Actiniaria Acontiaria sp 1

Cnidaria Anthozoa Actiniaria Actiniaria sp 1

Cnidaria Anthozoa Actiniaria Athenaria sp 2

Cnidaria Anthozoa Pennatulacea Halipteridae Halipteris sp 3

Cnidaria Anthozoa Pennatulacea Pennatulidae Ptilosarcus gurneyi 2

Cnidaria Anthozoa Pennatulacea Virgulariidae Stylatula elongata 17

Cnidaria Anthozoa Pennatulacea Virgulariidae Virgularia spp 2

Cnidaria Anthozoa Pennatulacea Virgulariidae Virgulariidae spp 3

Cnidaria Anthozoa Spirularia Cerianthidae Cerianthidae sp 6

Cnidaria Anthozoa Spirularia Cerianthidae Pachycerianthus 

fimbriatus

2

Cnidaria Anthozoa Spirularia Cerianthidae Pachycerianthus sp 25

Cnidaria Anthozoa Hexacorallia spp 1

Cnidaria Hydrozoa Anthoathecata Tubulariidae Ectopleura sp 1

Echinodermata Asteroidea Paxillosida Ctenodiscidae Ctenodiscus crispatus 3

Echinodermata Echinoidea Camarodonta Strongylocentrotidae Strongylocentrotus 

fragilis

2

Echinodermata Echinoidea Clypeasteroida Dendrasteridae Dendraster 

excentricus

219

Echinodermata Echinoidea Spatangoida Schizasteridae Brisaster latifrons 159

Echinodermata Holothuroidea Apodida Chiridotidae Chiridota spp 2

Echinodermata Holothuroidea Apodida Synaptidae Leptosynapta spp 8

Echinodermata Holothuroidea Dendrochirotida Cucumariidae Cucumaria piperata 1

Echinodermata Holothuroidea Dendrochirotida Cucumariidae Cucumariidae sp 1

Echinodermata Holothuroidea Dendrochirotida Phyllophoridae Pentamera populifera 1

Echinodermata Holothuroidea Dendrochirotida Phyllophoridae Pentamera 

pseudocalcigera

4
66

Phylum Class Subclass/Order Family Taxon Total 

Abund

Echinodermata Holothuroidea Dendrochirotida Phyllophoridae Pentamera rigida 2

Echinodermata Holothuroidea Dendrochirotida Phyllophoridae Pentamera spp 13

Echinodermata Holothuroidea Molpadida Caudinidae Paracaudina chilensis 27

Echinodermata Holothuroidea Molpadida Molpadiidae Molpadia intermedia 2

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphiodia 

occidentalis

41

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphiodia periercta 2

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphiodia spp 1

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphiodia urtica 364

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphioplus 

macraspis

133

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphioplus 

strongyloplax

230

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphipholis pugetana 2

Echinodermata Ophiuroidea Ophiurida Amphiuridae Amphiuridae spp 20

Echinodermata Ophiuroidea Ophiurida Ophiactidae Ophiopholis bakeri 3

Echinodermata Ophiuroidea Ophiurida Ophiuridae Ophiura lutkeni 68

Echinodermata Ophiuroidea Ophiurida Ophiuridae Ophiura spp 12

Echinodermata Ophiuroidea Ophiuroidea spp 10

Entoprocta Solitaria Barentsiidae Barentsia parva 1

Foraminifera Foraminifera sp Biserial type 1 2

Foraminifera Foraminifera sp Biserial type 2 1

Foraminifera Foraminifera sp Planispiral evolute type 1 34

Foraminifera Foraminifera sp Planispiral involute type 1 12

Foraminifera Foraminifera sp Planispiral involute type 2 41

Foraminifera Foraminifera sp Star type 1 1

Foraminifera Foraminifera sp Streptospiral type 1 1

Foraminifera Foraminifera sp Streptospiral type 2 1

Foraminifera Foraminifera sp Triserial type 1 250

Foraminifera Foraminifera sp Triserial type 2 17

Foraminifera Foraminifera sp Trochospiral type 1 24

Foraminifera Foraminifera sp Trochospiral type 2 7

Foraminifera Foraminifera sp Uniserial type 1 43

Foraminifera Foraminifera sp Uniserial type 2 52

Foraminifera Foraminifera sp Uniserial type 3 7

Foraminifera Foraminifera spp 3

Hemichordata Enteropneusta Enteropneusta Harrimaniidae Saccoglossus sp 1

Hemichordata Enteropneusta Enteropneusta Ptychoderidae Glossobalanus sp 5

Nematoda Nematoda spp 71

Nemertea Enopla Monostilifera Amphiporidae Amphiporus spp 4

Nemertea Enopla Monostilifera Emplectonematidae Paranemertes 

californica

3

Nemertea Enopla Monostilifera Tetrastemmatidae Tetrastemma spp 10

Nemertea Hoplonemertea Hoplonemertea spp 33

Nemertea Palaeonemertea Carinomidae Carinoma mutabilis 216
67

Phylum Class Subclass/Order Family Taxon Total 

Abund

Nemertea Palaeonemertea Tubulanidae Tubulanus 

polymorphus

51

Nemertea Palaeonemertea Tubulanidae Tubulanus spp 19

Nemertea Pilidiophora Heteronemertea Lineidae Cerebratulus 

marginatus

1

Nemertea Pilidiophora Heteronemertea Lineidae Cerebratulus 

montgomeryi

2

Nemertea Pilidiophora Heteronemertea Lineidae Cerebratulus spp 33

Nemertea Pilidiophora Heteronemertea Lineidae Lineidae spp 19

Nemertea Pilidiophora Heteronemertea Lineidae Lineus bilineatus 19

Nemertea Pilidiophora Heteronemertea Lineidae Lineus sp 4

Nemertea Pilidiophora Heteronemertea Lineidae Micrura spp 75

Nemertea Nemertea spp 3

Phoronida Phoronidae Phoronidae sp 118

Phoronida Phoronidae Phoronis spp 101

Platyhelminthes Rhabditophora Polycladida Leptoplanidae Leptoplanidae sp 6

Porifera Hexactinellid Hexactinellid sp 9

Porifera Porifera spp 5

Retaria Radiolaria Radiolarian spp 65

Sipuncula Sipunculidea Golfingiida Golfingiidae Golfingia spp 1

Sipuncula Sipunculidea Golfingiida Golfingiidae Golfingiidae sp 18

Sipuncula Sipunculidea Golfingiida Golfingiidae Nephasoma 

diaphanes

69

Sipuncula Sipunculidea Golfingiida Golfingiidae Nephasoma spp 11

Sipuncula Sipunculidea Golfingiida Golfingiidae Thysanocardia nigra 33

Sipuncula Sipunculidea Phascolosoma 

agassizii

2

Sipuncula Sipuncula spp 1

Sipuncula Sipunculida sp 1
68

Table A-6. All combinations of variables considered for the LINKTREE analysis.

Rows are sort by trees with the fewest number of splits at the top. The most parsimonious tree used all environmental 

factors with majority coverage (see Table 1 in Chapter 3). An equivalent tree was determined using only latitude, 

depth, median grain size, and total organic carbon (TOC). Since many sampling programs do not include analysis of 

TOC, and a tree with only two additional spits could be constructed without the use of that additional variable, we 

chose latitude, depth, and median grain size as the most parsimonious tree with the fewest variables (highlighted).

# Factors Variables Breaks to

all all BQ

4 latitude, depth, medianGS, TOC BQ

4 latitude, depth, medianGS, UpYear BR

4 latitude, depth, medianGS, pH BR

4 latitude, depth, medianGS, temp BR

3 latitude, depth, medianGS BS

4 latitude, depth, medianGS, UpMon BT

5 latitude, depth, medianGS, TOC, UpPrev BV

4 latitude, depth, medianGS, sal BW

4 latitude, depth, medianGS, DO BW

4 latitude, depth, fines, UpMon BX

4 latitude, depth, medianGS, UpPrev BX

4 latitude, depth, fines, UpYear BX

5 latitude, depth, medianGS, fines, upPrev BY

1 depth BY

4 latitude, depth, fines, UpPrev BY

4 latitude, depth, medianGS, fines BZ

3 latitude, depth, fines BZ

4 latitude, depth, fines, temp BZ

4 latitude, depth, fines, pH BZ

3 latitude, depth, salinity CA

4 latitude, depth, fines, DO CB

4 latitude, depth, fines, sal CB

3 depth, medianGS, upYear CC

3 latitude, depth, pH CC

3 depth, medianGS, UpMon CC

3 depth, medianGS, UpYear CC

1 fines CD

1 latitude CD

2 latitude, depth CD

3 latitude, depth, temp CE

3 latitude, depth, TOC CE

3 latitude, depth, DO CF

3 latitude, medianGS, DO CG

3 depth, medianGS, upPrev CI

3 depth, medianGS, temp CJ

3 depth, medianGS, DO CJ
69

# Factors Variables Breaks to

5 depth, fines, medianGS, temp, pH CM

3 latitude, medianGS, salinity CM

3 latitude, medianGS, pH CP

3 latitude, medianGS, temp CP

3 depth, medianGS, pH CQ

3 depth, medianGS, sal CQ

3 depth, medianGS, UpYear CR

2 latitude, medianGS CS

3 depth, medianGS, fines CU

2 depth, medianGS CW

3 depth, medianGS, temp CY

2 latitude, fines CZ

2 depth, fines DB

3 median grain size DF
70

Figure A-1. Correlations among all the environmental variables collected in the study.
71

Figure A-2. Total Organic Carbon (TOC) versus depth for the 256 stations (of 378 analyzed) for 

which we had TOC data.
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