APPENDIX AIR10-C Technical Data Reports Containing Habitat Maps at Local and Regional Scales

TDR MI-3 - Orange Sea Pens TDR

PORT METRO VANCOUVER | Roberts Bank Terminal 2 Information Request Response

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ROBERTS BANK TERMINAL 2 TECHNICAL DATA REPORT Marine Invertebrates Orange Sea Pens (Ptilosarcus gurneyi)

Prepared for: Port Metro Vancouver 100 The Pointe, 999 Canada Place Vancouver, BC V6C 3T4

Prepared by: Hemmera Envirochem Inc. 18th Floor, 4730 Kingsway Burnaby, BC V5H 0C6

And

Archipelago Marine Research Ltd. 525 Head Street Victoria, BC V9A 5S1

File: 302-042.02 November 2014

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens November 2014

Technical Report / Technical Data Report Disclaimer

The Canadian Environmental Assessment Agency determined the scope of the proposed Roberts Bank Terminal 2 Project (RBT2 or the Project) and the scope of the assessment in the Final Environmental Impact Statement Guidelines (EISG) issued January 7, 2014. The scope of the Project includes the project components and physical activities to be considered in the environmental assessment. The scope of the assessment includes the factors to be considered and the scope of those factors. The Environmental Impact Statement (EIS) has been prepared in accordance with the scope of the Project and the scope of the assessment specified in the EISG. For each component of the natural or human environment considered in the EIS, the geographic scope of the assessment depends on the extent of potential effects.

At the time supporting technical studies were initiated in 2011, with the objective of ensuring adequate information would be available to inform the environmental assessment of the Project, neither the scope of the Project nor the scope of the assessment had been determined.

Therefore, the scope of supporting studies may include physical activities that are not included in the scope of the Project as determined by the Agency. Similarly, the scope of supporting studies may also include spatial areas that are not expected to be affected by the Project.

This out-of-scope information is included in the Technical Report (TR)/Technical Data Report (TDR) for each study, but may not be considered in the assessment of potential effects of the Project unless relevant for understanding the context of those effects or to assessing potential cumulative effects. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - i - November 2014

EXECUTIVE SUMMARY

Port Metro Vancouver (PMV) is assessing the potential to develop the Roberts Bank Terminal 2 Project (RBT2 or the Project), a new three-berth marine terminal at Roberts Bank in Delta, B.C. The Project is part of PMV’s Container Capacity Improvement Program (CCIP), a long-term strategy to deliver projects to meet anticipated growth in demand for container capacity to 2030.

Hemmera has been retained by PMV to undertake environmental studies related to the Project. This technical data report focusses on the orange sea pen (Ptilosarcus gurneyi), a colonial octocoral that is broadly distributed along the Pacific coast of North America, from Alaska to California. A large aggregation, or bed, of orange sea pens, has been consistently observed at Roberts Bank, along the delta-front slope off the seaward face of Westshore Terminals (Golder 1996, Triton 2004, Archipelago 2009). The current state of knowledge on this species is limited, and many aspects of its biology and ecology remain poorly understood, both in B.C. waters and throughout the rest of its range. To address data gaps at both local and regional scales, this report is divided into three major sections: i) available literature and data review; ii) site-specific study describing the ecology and distribution of orange sea pens at Roberts Bank; and iii) local knowledge study compiling anecdotal ecological and biological information on orange sea pens across their geographic range.

Building on previous surveys conducted in 2003 and 2008, towed underwater video (SIMS) and SCUBA dive surveys were conducted in September 2011, with the objective to quantify orange sea pen distribution and densities at Roberts Bank, as well as document associations with other species, including fish and macroinvertebrates. Field data were supplemented by a local knowledge survey, conducted in January 2013, which collected anecdotal information on orange sea pen ecology, distribution, and value by the way of a questionnaire.

SIMS and SCUBA results at Roberts Bank suggest that: i) the spatial extent of orange sea pens is greater than what has been previously documented; ii) the orange sea pen bed is not comprised of a single age class, as was previously understood; iii) there appears to be a lack of natural predators; iv) orange sea pens provide habitat for a number of species; and v) other fauna (i.e., crustaceans, sea stars, anemones, and fish) are more likely to occur within areas of continuous to dense sea pens relative to areas where distribution is few to patchy or absent.

Forty-three responses (51% response rate) were received from the local knowledge survey, providing valuable information on geographical range, ecological significance, abiotic drivers, and value of orange sea pens in the northeast Pacific. Taken together, these responses provide a comprehensive snapshot of our current understanding of this poorly studied species. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - ii - November 2014

TABLE OF CONTENTS

EXECUTIVE SUMMARY ...... I 1.0 INTRODUCTION ...... 1

1.1 PROJECT BACKGROUND ...... 1

1.2 ORANGE SEA PEN STUDY OVERVIEW ...... 1 2.0 REVIEW OF AVAILABLE LITERATURE AND DATA ...... 3

2.1 TAXONOMY AND DISTRIBUTION ...... 3

2.2 LIFE HISTORY AND BEHAVIOUR ...... 4

2.3 ECOLOGICAL ROLE ...... 5 2.3.1 Trophic Interactions ...... 5 2.3.2 Species Associations ...... 5 2.3.3 Habitat Value ...... 6

2.4 HABITAT REQUIREMENTS AND LIMITING FACTORS ...... 7

2.5 CONSERVATION STATUS ...... 8

2.6 SUMMARY OF PREVIOUS STUDIES AT ROBERTS BANK ...... 8 3.0 ECOLOGICAL STUDY ...... 9

3.1 STUDY AREA ...... 9

3.2 TEMPORAL SCOPE...... 9

3.3 STUDY METHODS ...... 9 3.3.1 SIMS Survey Methods & Data Analysis ...... 9 3.3.2 SCUBA Survey Methods and Data Analysis ...... 15

3.4 RESULTS ...... 15 3.4.1 SIMS Results ...... 15 3.4.1.1 Delta Front Slope ...... 15 3.4.1.2 Existing Dredge Basin ...... 17 3.4.1.3 Roberts Bank Overall ...... 17 3.4.2 SCUBA Results ...... 25 3.4.2.1 Dive Site SP1 ...... 25 3.4.2.2 Dive Site SP2 ...... 25 3.4.2.3 Dive Site SP3 ...... 26

3.5 KEY FINDINGS ...... 27 3.5.1 Discussion of Key Findings ...... 27 Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - iii - November 2014

3.5.2 Data Gaps and Limitations ...... 30 4.0 LOCAL KNOWLEDGE STUDY ...... 31

4.1 STUDY METHODS ...... 31

4.2 RESULTS ...... 32 4.2.1 Geographical Range and Ecological Significance ...... 32 4.2.1.1 Species Distribution ...... 32 4.2.1.2 Observed Densities ...... 32 4.2.1.3 Ecologically-Linked Species ...... 32 4.2.2 Abiotic Drivers ...... 33 4.2.2.1 Substrate Type ...... 33 4.2.2.2 Flow ...... 33 4.2.2.3 Depth ...... 33 4.2.2.4 Unique Habitat Attributes ...... 33 4.2.2.5 Significance of Abiotic Factors ...... 36 4.2.2.6 Additional Abiotic Factors ...... 36 4.2.3 Ascribed Value ...... 36 4.2.3.1 Ecosystem Functions ...... 36 4.2.3.2 Anthropogenic Value ...... 36 4.2.3.3 Additional Comments ...... 37 5.0 CLOSURE ...... 38 6.0 REFERENCES ...... 39 7.0 STATEMENT OF LIMITATIONS ...... 44

List of Tables

Table 1-1 Orange Sea Pen Study Components and Major Objectives ...... 1 Table 3-1 Faunal Distribution Class Codes for SIMS Bio-classification ...... 11 Table 3-2 Summary of the Sea Pen Dive Survey Conducted on Roberts Bank October 19, 2011 .. 15 Table 3-3 Summary of Predictive Diagnostic Coefficients in the Best-fit Generalised Linear Models for Four Faunal Groups at Roberts Bank ...... 17 Table 3-4 Sea Pen Density Data for Dive Sites SP2 (sparse) and SP3 (dense) at Roberts Bank on October 19, 2011 ...... 26 Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - iv - November 2014

List of Figures

Figure 3-1 Orange Sea Pen Study Area at Roberts Bank ...... 12 Figure 3-2 Towed Underwater Video (SIMS) Tracklines over Time (2003, 2008, 2011) at Roberts Bank ...... 13 Figure 3-3 2011 Towed Underwater Video (SIMS) Tracklines and SCUBA Dive Sites at Roberts Bank ...... 14 Figure 3-4 Orange Sea Pen Distribution at Roberts Bank ...... 19 Figure 3-5 Slope Profiles along Orange Sea Pen Density Gradients ...... 20 Figure 3-6 Cumulative Distribution of Crustaceans from SIMS Surveys (2003, 2008, 2011) at Roberts Bank ...... 21 Figure 3-7 Cumulative Distribution of Sea Stars from SIMS Surveys (2003, 2008, 2011) at Roberts Bank ...... 22 Figure 3-8 Cumulative Distribution of Anemones, Sea Cucumbers, and Snails from SIMS Surveys (2003, 2008, 2011) at Roberts Bank ...... 23 Figure 3-9 Cumulative Distribution of Marine Fishes from SIMS Surveys (2003, 2008, 2011) at Roberts Bank ...... 24 Figure 4-1 Anecdotal Sea Pen Observations along the Pacific Coast ...... 34 Figure 4-2 Anecdotal Sea Pen Observations in the Strait of Georgia ...... 35

List of Appendices

Appendix A Photographs

Appendix B Statistical Summaries

Appendix C Local Knowledge Survey Questionnaire and Results

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 1 - November 2014

1.0 INTRODUCTION

This section provides an overview of the study, including project background and the study components and major objectives.

1.1 PROJECT BACKGROUND

The Roberts Bank Terminal 2 Project (RBT2 or Project) is a proposed new three-berth marine terminal at Roberts Bank in Delta, B.C. that could provide 2.4 million TEUs (twenty-foot equivalent unit containers) of additional container capacity annually. The Project is part of Port Metro Vancouver’s Container Capacity Improvement Program, a long-term strategy to deliver projects to meet anticipated growth in demand for container capacity to 2030.

This technical data report (TDR) describes the results of orange sea pen (Ptilosarcus gurneyi) studies and is split into two sections: i) ecological survey; and ii) local knowledge survey. Methods, results, and discussion are described below for each study.

1.2 ORANGE SEA PEN STUDY OVERVIEW

A review of available information and state of knowledge was completed for orange sea pens to identify key data gaps and areas of uncertainty within the general Project area. This TDR describes the study findings for key components identified from this gap analysis. Study components with their major objectives and a brief overview are provided in Table 1-1.

Table 1-1 Orange Sea Pen Study Components and Major Objectives

Component Major Objectives Brief Overview A towed underwater video (SIMS) survey was SIMS (Subtidal Imaging Map and classify the geographic conducted in 2011 to map their spatial extent at and Mapping System) extent of sea pen distribution at Roberts Bank. Builds on SIMS surveys survey Roberts Bank. conducted in the same general area of Roberts Bank in 2003 and 2008. SCUBA surveys were conducted in 2008 and 2011 at three sites along a sea pen density Collect sea pen density and SCUBA survey gradient to quantify densities and to associated species information. characterise the associated biophysical environment. Gain a better understanding of the Individuals possessing relevant expertise, Local Knowledge spatial extent, life history, and experience, and/or knowledge pertaining to Survey ecological importance of sea pens orange sea pens were contacted for their input along the northeast Pacific coast. via a local knowledge questionnaire. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 2 - November 2014

The current state of knowledge on orange sea pens, and on cold-water corals in general, is limited. Little is known about their abundance, distribution, or ecological role in B.C. and throughout the rest of their geographic range. Many aspects of orange sea pen biology and ecology are not well studied or understood, and often based on speculation about possible similar traits with species in other regions, such as the Arctic, North Atlantic, and tropical environments.

The ecological study component builds on previous work conducted by Archipelago (Triton 2004, Archipelago 2009) using SIMS and SCUBA surveys to characterise orange sea pen spatial extent and densities, and attempts to better understand ecosystem interactions of orange sea pens, and the extent to which dense aggregations may provide habitat complexity and structural relief for macroinvertebrates and fish. The local knowledge survey addresses several data gaps around orange sea pen biology and ecology through the collection of anecdotal information.

Data on abiotic drivers of orange sea pen density and distribution are presented and discussed in the Habitat Suitability Modelling TDR (Hemmera 2014a), which was used to map areas of optimal sea pen habitat at Roberts Bank. Additionally, the Benthic Subtidal TDR (Hemmera 2014b) provides additional data on orange sea pen densities, habitat use, and ecosystem interactions at depth ranges (up to -40 m CD), which were beyond the limit of SIMS.

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 3 - November 2014

2.0 REVIEW OF AVAILABLE LITERATURE AND DATA

A thorough review was undertaken for information on orange sea pens, specifically their ecology and habitat preferences at Roberts Bank and across their entire geographic range. Numerous literature and data sources were consulted, including:

 Publicly available Aboriginal Traditional Knowledge (ATK);

 Workshops with Tsawwassen First Nation (TFN);

 Books;

 Academic journals;

 Databases (e.g., DFO WAVES Catalogue; NOAA);

 Consultant reports;

 Government technical reports; and

 Expert opinion.

No ATK references to orange sea pens were identified and TFN community members indicated that the species holds no cultural or socioeconomic value to their community. Less than ten studies on orange sea pen were found, reflecting the paucity of literature and how cursory our understanding of this species remains. Given the limited information, this report will be informed by work on related species and in other areas (e.g., Arctic, Atlantic, tropics).

2.1 TAXONOMY AND DISTRIBUTION

Sea pens are octocorals in the order Pennatulacea and, along with the true soft corals and gorgonians, form the subclass Octocorallia within class Anthozoa and phylum Cnidaria. There are 16 families of sea pens with approximately 300 species occupying both tropical and temperate waters worldwide. According to the World Register of Marine Species (WoRMS 2008), the genus Ptilosarcus is comprised of two species, including the orange sea pen. In the Pacific Northwest, orange sea pens appear to be the most common sea pen in shallow water, while several other species of sea pens (Virgularia sp., Stylatula sp. and Acanthoptilum gracile) have also been documented (Lamb and Hanby 2005).

Individual sea pens are actually colonies of many polyps (like a coral head) rather than a single animal (like an anemone), and the polyps show an eight-fold symmetry (Fuller et al. 2008). Orange sea pens, and pennatulaceans in general, are known to form dense aggregations, known as sea pen beds, but are otherwise broadly distributed at low density (Fuller et al. 2008). Individual colonies can consist of tens of thousands of polyps, and a large sea pen bed can easily comprise over 40,000 individuals (Erhardt and Moosleitner 1998). Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 4 - November 2014

Orange sea pens are found throughout the northeastern Pacific from Alaska to Southern California (Shimek 2011), and at a depth range that includes the lowest intertidal zone to depths of about 150 metres, but they are most abundant in shallow waters (Shimek 2011). In the 1960s, Birkeland (1969) documented an extensive zone of dense orange sea pens (up to 22/m2) in depths of -10 to -25 m from Olympia to Everett Washington, a distance of over 150 km of coastline. Density was lower at both shallower (0 to -10 m) and deeper (-25 to -50m) depths. A more recent study by Kyte (2001) showed that the large populations described by Birkeland (1969) are no longer present and remaining populations are relatively sparse and patchy.

2.2 LIFE HISTORY AND BEHAVIOUR

Sea pens are sessile (i.e., immobile) macroinvertebrates, living in unconsolidated ocean bottom sediments. They are not fastened to the substrate and are capable of locomotion by crawling out of the sediment, inflating with water, and drifting in the currents. Fully-expanded, large adult individuals may extend 60 cm above the seafloor, with their base (termed “peduncle”) burrowed 15 to 30 cm into the sediment (Shimek 2011). Orange sea pens may live up to 15 years and take five or six years to reach sexual maturity (Birkeland 1974).

In orange sea pens, the sexes are separate (i.e., each colony of polyps is either male or female), and reproduction is sexual, through broadcast spawning of gametes (Edwards and Moore 2008). Eggs and sperm are released through the mouths of the polyps and fertilization takes place externally (Chia and Crawford 1973). The total fecundity of female sea pens is high, ranging from approximately 30,000 to 200,000 eggs per colony (Chia and Crawford 1973, Soong 2005). In B.C. waters, orange sea pens spawn in late March, generally in the first week following the spring equinox (Chia and Crawford 1973, Shimek 2011). The free-swimming larvae do not feed and will settle within seven days if a suitable substratum, such as coarse sand, is encountered (Chia and Crawford 1973).

Studies in Puget Sound by Birkeland (1968, 1974) indicate that larval settlement can be patchy in space and highly episodic in time giving rise to discontinuous populations differing in age and size. Large year- to-year differences in recruitment rates were also seen in Renilla kollikeri, a sea pen from the coast of California (Davis and VanBlaricom 1978). Chia and Crawford (1973) posit that the nature of the substratum at the time of larval settlement dictates recruitment success in any given area, and hence the patchiness of distribution. They further suggest that stochastic recruitment patterns of orange sea pens, in both time and space, are actually a major defense mechanism, making the species generally unavailable to predators. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 5 - November 2014

Orange sea pens can contract their bodies and burrow into the sediment by expelling water and mucus from their hydroskeleton (Kozloff 1993). They alternately expand for feeding and contract into the sediment at irregular intervals, a behavioural pattern apparently unrelated to environmental factors such as current velocity, turbidity, and light level (Birkeland 1974, Dickinson 1978) or available food supply (Shimek 2011). Although the ecological significance of this behavioural rhythm is uncertain, burrowing may allow orange sea pens to be less conspicuous to predators (Birkeland 1974). While diving in Puget Sound, Birkeland (1968) observed that only about 26% of the sea pens were exposed at any one time.

2.3 ECOLOGICAL ROLE

2.3.1 Trophic Interactions

Orange sea pens are passive suspension feeders whose diet consists mainly of phytoplankton (Best 1988). In turn, they are fed upon by several species of predatory sea stars and nudibranchs. Of the nudibranch predators, striped (Armina californica) and diamond back (Tritonia festiva) nudibranchs feed exclusively on sea pens, while the opalescent nudibranch (Hermissenda crassicornis) has a relatively diverse diet, and feeds on a variety of other invertebrates as well (Birkeland 1974). Adult sea pens are mostly eaten by sea stars (Birkeland 1974). The spiny red sea star (Hippasteria spinosa) is a specialised orange sea pen predator, while rose (Crossaster papposus) and vermillion (Mediaster aequalis) stars consume sea pens opportunistically. Leather stars (Dermasterias imbricata) consume a variety of prey sources, but when found in sea pen beds, feed almost exclusively on them. Birkeland’s studies (1969, 1974) describe the orange sea pen’s ecological importance as being keystone, particularly for the predators that feed exclusively on them; however, there are no reports of fish or macroinvertebrates (e.g., Dungeness crabs, Metacarcinus magister) feeding directly on orange sea pens, and accounts of predation on other sea pen species are limited.

2.3.2 Species Associations

Habitat complexity and heterogeneity have been linked to changes in organism abundance and diversity in a variety of terrestrial and aquatic settings (Bell 1985, Levin and Dayton 2009). Sea pen beds provide important structure in low-relief sand and mud habitats where there is little physical habitat complexity, thus providing shelter from currents and predators (Tissot et al. 2006). Sea pens are also considered “ecosystem engineers”, which shape the environment by burying into soft sediments, enabling deep oxygen penetration as well as modifying hydrodynamics, and allowing nutrients and plankton to be retained near the sediment thereby supplying the detrital food chain (Tissot et al. 2006, Boutillier et al. 2010). Sea pens are thought to contribute substantially to the species richness of their respective environments, particularly in terms of small planktonic and benthic invertebrates that, in turn, may be preyed upon by fishes (Hughes 1998, Beaulieu 2001, Buhl-Mortensen and Mortensen 2004, Tissot et al. 2006). Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 6 - November 2014

Sea pens may provide habitat complexity, and structural relief for fish and macroinvertebrates that favour emergent structures (e.g., juvenile flatfish) (Ryer et al. 2004, 2007, Pirtle 2005, Stoner et al. 2007). In Alaska, cods and pollocks (family Gadidae) were frequently caught with sea whips and sea pens, including orange sea pens, as bycatch (Malecha et al. 2005). Fisherman from Port-Aux-Basques, Newfoundland identified areas with sea pens as being good fishing grounds, claiming areas with sea pens have more and larger Atlantic cod and halibut than areas without (Colpron et al. 2010). Stripetail rockfish (Sebastes saxicola) were observed using sea pens (primarily Stylatula spp.) as a nursery ground in California (Field et al. 2001), and locally in Puget Sound, the density of geoducks (Panopea generosa) was significantly correlated with orange sea pens (Goodwin and Pease 1991). Several of these documented associations were hypothesised to be coincidental (e.g., rockfish, geoduck), whereby fish or macroinvertebrates share similar habitat requirements with sea pens (e.g., in areas of high flows for enhanced prey delivery), but have no direct association or functional relationship (Auster 2005).

Other studies have not found relationships between fish distribution and sea pens (Tissot et al. 2006, Edinger et al. 2007, Hixon and Tissot 2007). While results indicated the fish and invertebrates considered in these studies did not have a strong or obligate relationship, the authors suggest that the importance of soft corals, sea pens, and small gorgonians as potential fish and invertebrate habitat, particularly for juveniles, should not be overlooked.

2.3.3 Habitat Value

Considering the knowledge gaps that exist for orange sea pens, they likely provide a number of ecosystem functions that are not yet fully understood (Diaz et al. 2003). The value of sea pen aggregations in general is increasingly being recognised by international organizations and governments in parts of Canada, the United States, and Europe.

In the Gulf of St. Lawrence, Fisheries and Oceans Canada (DFO) considers high concentrations of sea pens and sponges as habitats suitable for the establishment of highly diverse benthic communities, and recognises sea pens as important habitat for both fish and invertebrates (DFO 2012). Concern over destructive fishing methods, such as bottom trawling, has led corals (including sea pens) to be defined as “sensitive” and “unique” habitats within the Laurentian Channel Area of Interest (AOI) (DFO 2012). In the Canadian Arctic, Northern Baffin Bay was designated as a new Ecologically and Biologically Significant Area (EBSA) based on the presence of significant concentrations of sea pens (Ombellula sp.) (Kenchington et al. 2011). More locally, the report, Ecosystem Overview Report of Pacific North Coast Management Area (PNCIMA) by DFO (2007), highlighted that cold-water corals (including unspecified species of sea pens) and sponges may provide essential habitat for some benthic organisms, including many juvenile and some adult fish species (especially rockfish). Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 7 - November 2014

In Alaska, sea pens, sea whips, corals, sponges, and other “living substrates” have been identified as Habitat Areas of Particular Concern (HAPC) (Heifetz 2002). A HAPC is defined as a habitat that is ecologically important, sensitive to disturbance, stressed, or rare. HAPCs are not given any regulatory protection; however, projects with potential adverse impacts to HAPCs tend to be more heavily scrutinised during the assessment process (NOAA 2011). Further, in New England, sea pens, stony corals, and soft corals/gorgonians were identified as structural components of fish habitat in a vulnerability assessment, conducted as part of a model to evaluate the impacts of fishing on essential fish habitat (NOAA 2010).

The United Nations Food and Agriculture Organization (FAO) recognises certain cold-water corals, including representatives from class Octocorallia, to which sea pens belong, as meriting vulnerable marine ecosystem (VME) status given their vulnerability to most kinds of bottom fishing (FAO 2008). The Northwest Atlantic Fisheries Organization (NAFO) also identified sea pen beds as a component of Vulnerable Marine Ecosystems, while simultaneously acknowledging that not all corals and sponges meet criteria associated with vulnerability (Fuller et al. 2008). Sea pens are also included on an initial OSPAR1 list of Threatened and/or Declining Species and Habitats, recognising the role sea pens play in creating complex habitats with higher macrofaunal species diversity (OSPAR 2010).

2.4 HABITAT REQUIREMENTS AND LIMITING FACTORS

According to a review of sub-tidal benthic habitats and invertebrate biota in the Strait of Georgia, orange sea pens are characteristic of shallow (0 to 30 m) silt/sand habitats (Burd et al. 2008). The size distribution of particles in marine sediment is an important ecological parameter that can have a major influence on the composition of the biological community (Hughes 1998). While sea pens are anchored within the sediment, they do not depend upon it for food; however, particle size and associated organic content are thought to be the two major factors inducing orange sea pen larval settlement (Chia and Crawford 1973). If appropriate sediments are not available, settlement and metamorphosis may be delayed for as much as a month, and without appropriate sediment, the sea pen will die (Shimek 2011).

As passive suspension feeders, orange sea pens are highly dependent on ambient flow conditions for feeding (Best 1988), but are also capable of shaping the environment by modifying local hydrodynamics to optimise their food uptake (Buhl-Mortensen et al. 2010). Burd et al. (2008) reported finding sea pens and sea whips in higher current areas. Through a series of laboratory experiments, Best (1988) demonstrated that sea pen volume flow rates (i.e., water that passes through a sea pen) initially increase with increasing flow speed, but subsequently peak and decline because the organism becomes so bent back by the flow that more water tends to flow over, rather than through, the filter. These findings suggest that there is an optimal flow range wherein sea pens are able to maximise access to food without being physically deformed, or uprooted, by the flow.

1 The OSPAR Convention is the current legal instrument guiding international cooperation on the protection of the marine environment of the Northeast Atlantic. Work under the Convention is managed by the OSPAR Commission, made up of representatives of the Governments of 15 Contracting Parties and the European Commission, representing the European Union. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 8 - November 2014

2.5 CONSERVATION STATUS

Orange sea pens are not protected under any provincial or federal legislation.

In 2011, DFO released a Pacific Region Cold-Water Coral and Sponge Conservation Strategy that recognises the sensitivity of these biogenic habitats to human activities, and aims to promote their conservation, health and integrity (DFO 2011). Orange sea pens are one of 80 species of cold-water corals and sponges covered under this strategy (J. Finney, Fisheries and Oceans Canada, personal communication).

2.6 SUMMARY OF PREVIOUS STUDIES AT ROBERTS BANK

As described in Archipelago (2009), studies of the subtidal environment at Roberts Bank spanning the last two decades have included observations of the orange sea pen, particularly in the vicinity of the northwest corner of Westshore Terminals (Gartner Lee 1992, Golder 1996, Triton 2004).

A large orange sea pen bed was first delineated and mapped in September 2003 by Archipelago (Triton 2004) using a SIMS, or towed underwater video system. Survey results reported a continuous to densely distributed bed of sea pens covering approximately 15 hectares in the sandy substrate northwest of Westshore Terminals, from -2.5 to -18 m chart datum (CD) (Triton 2004). A larger area with few to patchy distribution of sea pens, covering an area of approximately 66 ha, surrounded the dense bed and extended from -2 to -24 m CD (Triton 2004). Dungeness crab, spiny pink stars (Pisaster brevispinus), and starry flounder (Platichthys stellatus) were common in the area.

In 2008, Archipelago conducted a follow-up SIMS survey to confirm the sea pen polygons previously mapped in Triton (2004), and to extend the survey boundaries. Results indicated no change in the boundaries or depth range of the dense sea pen bed, but indicated a larger area (approximately 114 ha) of few to patchy sea pens, extending to -35 m CD. Dungeness crabs and flatfish (including starry flounder) were commonly observed in the area, as were spiny dogfish (Squalus suckleyi) (Archipelago 2009).

Following the SIMS survey in 2008, Archipelago also conducted a SCUBA survey to collect sea pen density and community assemblage information. Sea pen density ranged from 0 to 1/m2 (mean= 0.08/m2) in the patchy area, and from 1 to 8/m2 (mean= 4.3 /m2) in the denser area (Archipelago 2009).The dense site was characterised by silty-sand with 25 to 50% cover of diatoms. Other species associated with the sea pens at the dense site included, tube dwelling worms, juvenile Dungeness crabs, and flatfish (predominantly speckled sanddab, Citharichthys sordidus). The patchy site was characterised by silty- sand with <25% cover of diatoms and associated species included, tube worms, several juvenile crabs, a sunflower star (Pycnopodia helianthoides), spiny pink stars, and flatfish (predominantly speckled sanddab). The site with no sea pens was also comprised of silty-sand with some areas of exposed clay and 25 to 50% cover of diatoms and other observed species included, tube worms, Dungeness crabs, and flatfish. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 9 - November 2014

3.0 ECOLOGICAL STUDY

The ecological study, conducted in 2011, has two major components:

(i) SIMS Survey; and (ii) SCUBA Survey.

3.1 STUDY AREA

The study area encompassed areas of the Roberts Bank delta front slope, from the BC Ferries Terminal in the south to Canoe Passage in the north, including the existing Deltaport dredge basin in the Inter- causeway Area (Figure 3-1). Minimum depth was the 0 m contour, as orange sea pens are a subtidal species, while maximum depth was limited by constraints of the SIMS system to approximately -35 m CD. Sampling effort was highest in the vicinity of the proposed Project as this area houses a large continuous to densely distributed area of orange sea pens; however, boundaries were expanded to delineate and map the spatial extent of orange sea pens at Roberts Bank in order to gain a better understanding of the sub-regional context of this species’ distribution.

3.2 TEMPORAL SCOPE

Sea pens are capable of locomotion such that aggregations are not predictable and have the potential to show appreciable variation over time. In consideration of such temporal variability, the scope of orange sea pen studies intends to capture baseline conditions in the study area. Studies build on work dating back to 2003, and were conducted over a two-year period from 2011 to 2013.

3.3 STUDY METHODS

3.3.1 SIMS Survey Methods & Data Analysis

Archipelago Marine Research Ltd. performed an underwater video survey using SIMS from September 15 to 17, 2011. SIMS is a towed video system developed to carry out systematic mapping of marine vegetation, macroinvertebrates, seafloor substrates, and morphology from the intertidal zone to depths of approximately -35 m CD.

The SIMS survey builds on previous SIMS work conducted in the area, with tracklines positioned through the sea pen polygons mapped in 2003 and 2008 to confirm that sea pens were still present at the documented locations and in similar densities (Figure 3-2). A grid spacing of 100 x 100 m was used in the existing sea pen area, and the boundaries were extended approximately 1 km in each direction to the northwest and southeast. A larger grid size (spacing between 250 m and 750 m) was then used to collect imagery within the existing dredge basin to 1 km southeast of the BC Ferries Terminal and 5 km northwest to Canoe Passage (Figure 3-3). The field of view was approximately 1 x 3 m with the camera maintained at an elevation of 1 to 1.5 m above the seafloor and towed at a speed of about 1 knot Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 10 - November 2014

(2 km/hr). The survey generated 47.7 km of video survey track lines, 19 hours of video imagery, and ranged in depth from -1.0 to -25 m CD. The depth of previous surveys extended to -35 m CD, which is reflected in several figures in this report.

Video imagery was classified by an Archipelago biologist in Victoria, B.C., for marine vegetation and fauna (i.e., crustaceans, anemones, sea stars, and fish) using a standard and biotic classification system initially developed for the Province of British Columbia (Harper et al. 1998a, b, 1999). The SIMS database system allows data entry for each second of video imagery collected, permitting the classifier to enter data every second if biotic features changed.

Orange sea pen distribution was classified according to the codes presented in Table 3-1, and the interpreted data were imported into ArcView for map production. Maps include point data and polygon data. Polygon data were determined through visual extrapolation of point data for particular biological features (e.g., sea pens). As SIMS faunal data is distribution data, densities, as number of organisms per square area, were not quantified from the SIMS imagery. Other fauna, including crustaceans, sea stars, anemones, and fish were also classified and mapped.

The categorical distribution data obtained from the SIMS surveys were used to model the influence of sea pen distribution, depth, and sample year on the distribution of four other faunal groups (crustacean, sea star, anemones, and fishes) using logistic regression. Four dependent variables, consisting of presence- absence data for each of the four faunal groups, were analysed separately using the following procedure. Categorical distribution data from SIMS were converted to presence-absence data for each faunal group, since distribution codes greater than 1 (“few”; Table 3-1) were rarely recorded. To maintain balance for model analyses, a commensurate number of absence points were randomly selected to balance each faunal group presence numbers.

The independent variable, sea pen habitat, was derived from SIMS distribution classifications by converting to three ordinal categories of sea pen density (i.e., outside the bed, few-patchy distribution, or continuous-dense distribution), corresponding to the SIMS categories No Observed Fauna, codes 1 and 2 combined, and codes 4 and 5 combined, respectively. While the SIMS surveys did not directly quantify densities of organisms, the correspondence of the three categories chosen for the logistic regression to an increasing trend in sea pen densities was confirmed by SCUBA observations (see Section 3.4.2). Data associated with the SIMS code 3, “uniform” distribution of sea pens, were excluded from the analysis as these observations were deemed not to be contributing information to densities but only to distribution of individuals, and their relationship to density had not been investigated by divers. The magnitude of difference among sea pen categories is not known, and thus the three density categories used for sea pen habitat were included in the regression model using a flexible step function. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 11 - November 2014

Of the other two independent variables included in the regression model, depth (m relative to CD) was included as a continuous variable, and sample year (2003, 2008, 2011, and 2012) as a categorical nominal variable. Individual models for each faunal group were selected based on Akaike’s Information Criterion (AIC). The best fit models had the fewest parameters within two AIC points of the minimum AIC score (Burnham and Anderson 2002). Estimation of GLM parameters and model selection were performed using the statistical package R.

Table 3-1 Faunal Distribution Class Codes for SIMS Bio-classification

Code Descriptor Distribution Example

NOF No fauna observed

A rare (single) or a few sporadic 1 Few individuals

A single patch, several 2 Patchy individuals, or a few patches

3 Uniform Continuous uniform occurrence

Continuous occurrence with a 4 Continuous few gaps

5 Dense Continuous dense occurrence

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Figure 3-1 Orange Sea Pen Study Area at Roberts Bank

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Figure 3-2 Towed Underwater Video (SIMS) Tracklines over Time (2003, 2008, 2011) at Roberts Bank

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Figure 3-3 2011 Towed Underwater Video (SIMS) Tracklines and SCUBA Dive Sites at Roberts Bank This page is intentionally left blank. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 15 - November 2014

3.3.2 SCUBA Survey Methods and Data Analysis

Divers determined orange sea pen density (numbers/m2) and community assemblage information within three reference areas: i) ‘continuous to dense’ sea pen distribution (SP3); ii) ‘few to patchy’ sea pen distribution (SP2); and, iii) ‘absent’ sea pens (SP1) (see Figure 3-2; Table 3-2), which were the same locations surveyed in 2008.

At each dive site, divers established two 6 x 2 m survey grids and used 1 m2 quadrats to count sea pen density. Six quadrats were counted per grid (i.e., every other square meter was sampled) for a total of 12 quadrats per site. Grids were approximately 3 m apart, and divers worked away from each other such that the greatest distance of separation was approximately 7 m. A greater distance of separation was not possible due to reduced visibility and the Work Safe BC regulatory requirement for the two divers to be in visual contact. At all three survey locations, biophysical features were documented to provide community assemblage information. Approximately 15 minutes of underwater video imagery were collected at each survey location.

Mean densities and community assemblage information are qualitatively described and compared among the three dive sites.

Table 3-2 Summary of the Sea Pen Dive Survey Conducted on Roberts Bank October 19, 2011

Dive Coordinates Tide Range Depth (m Survey Location Description Time Site (UTM) (m CD) CD)

West end of SIMS fine survey grid N5429593 12:26 to SP1 where sea pens were not encountered 4.1 3.5 in previous survey (2008). E485730 12:38 East end of SIMS fine survey grid where N5428978 10:25 to SP2 3.8 to 3.9 5 sea pen distribution is few to patchy. E487558 10:58

Center of SIMS fine survey grid where N5429162 11:30 to SP3 sea pen distribution is continuous to 4.0 to 4.1 3.5 dense. E486869 11:59 Note: Depths are reported relative to chart datum.

3.4 RESULTS

3.4.1 SIMS Results

3.4.1.1 Delta Front Slope

Documented orange sea pen habitat at Roberts Bank starts in the Inter-causeway Area and extends 3.5 km northwest, along the delta slope, at depths between -1.5 and -35 m CD. Sea pens are most prevalent in sandy substrate at depths less than -5 m CD. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 16 - November 2014

Surveys conducted in 2003 and 2008 identified the presence of a large (15 ha) area of continuous to densely distributed sea pens ranging from -2.5 to -18 m depth CD, which was confirmed in the 2011 survey. Additionally, a second dense aggregation was found on the 2011 survey off the southeast corner of Westshore Terminals, occupying an area of approximately 7.6 ha, and ranging from -3 to -19 m depth CD. Combined survey results across years at Roberts Bank yielded 23 ha of densely distributed sea pens surrounded by 151 ha of sparsely distributed sea pens (Figure 3-4). Sea pens were observed outside the delineated polygons, but not in high enough densities to be included within the distribution polygons, accounting for the data points present in Figure 3-4. Photographs are presented in Appendix A.

Sea pens were also documented between -2 to -18 m depth CD in the Inter-causeway Area (including the existing dredge basin), and east of the BC Ferries Terminal, where trackline spacing was coarser. The distribution of sea pens was few to patchy within the existing dredge basin east to the survey area boundary, with some observations of continuous to dense distributions within the Inter-causeway Area at depths between -2 to -10 m CD. Of note is that mapped distributions are a function of sampling effort (i.e., trackline spacing and length) and interpolation based on researcher experience and professional judgement. Additional and/or denser sea pen beds may be present in the area, but were not captured in this study.

Tracklines were also extended towards Canoe Passage to determine if orange sea pen distribution reaches further northwest; however, only one sea pen was observed outside the existing polygon in this direction, indicating limited suitable habitat. Relatively large sand waves and slumped sediment were documented, suggesting that the substrate may be too unstable for sea pen colonization. An active commercial crab fishery within documented sea pen areas may also have been a factor in distribution (Appendix A: Photo 5).

Slope profiles were created along sea pen density gradients (i.e., dense, sparse, absent) to identify areas along the delta front slope where sea pens tend to congregate (Figure 3-5). The slopes through the dense portion of the sea pen field were similar for both transects A and B, at m = 0.051 and m = 0.056, respectively. Sea pens were also sparse or absent along flat slopes (m= 0) and steeper slopes (m> 0.1) on both transects

Macroinvertebrate species commonly observed within orange sea pen areas included: Dungeness crab, sunflower stars; spiny pink stars; and plumose anemones (Metridium giganteum) (Figures 3-6 to 3-8). Flatfish (adult and juvenile), including starry flounder and Pacific sanddab (Citharichthys sordidus), were also commonly observed, while lingcod (Ophiodon elongatus) and kelp greenling (Hexagrammos decagrammus) were common on vegetated artificial rock reefs within the sea pen beds off the southern face of Westshore Terminals (Figure 3-9). All species described in the 2011 survey were also observed in 2003 and/or 2008 surveys. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 17 - November 2014

3.4.1.2 Existing Dredge Basin

Scattered orange sea pens, few to patchy in distribution, extend into the deepest portion of the existing dredge basin (-13 to -22 m depth CD). Additionally, they were observed in the northeastern segment of the existing dredge basin, where they were not documented in 2003; however, the extent of the surveys was not the same between years so a direct comparison can not be made. Orange sea pens appeared to be predominantly associated with sandy substrate, although some individuals were found in muddier substrate.

Within the existing dredge basin, orange sea pens and plumose anemones were the only epibenthic macroinvertebrate species documented. Mobile macroinvertebrate observations included: scattered Dungeness crabs; sunflower stars; spiny mud stars (Luidia foliolata); and mottled sea stars (Evasterias troschelli). Additionally, pandalid shrimp (Pandalus sp.) and a few snails were noted in the shallower portion of the dredge area, adjacent to the rip-rap of the existing Roberts Bank terminals. A number of fish were also noted in the rip-rap areas, including lingcod. Bacterial mats (Beggiatoa spp.) were found in the shallower portions of the dredged area, indicating some hypoxia in the sediments. Native eelgrass (Zostera marina) was the dominant vegetation observed on the shallowest portions of the mud/sand flats. All macroinvertebrates, fish, and vegetation described above, with the exception of the orange sea pens, were also documented in the 2003 SIMS survey.

3.4.1.3 Roberts Bank Overall

A best-fit GLM was selected from a set of models for each faunal group. Table 3-3 summarises the diagnostic coefficients for each model, while Appendix B presents complete results from the analyses. Orange sea pen habitat was present in best fit models for all four faunal groups. Depth and sampling year were included in the best fit models for crustaceans, anemones, and sea stars. Year was not included in the best fit model for fishes as there were only data for a single year, 2003. Correlation coefficients indicate the direction of relationship between likelihood of recording presence of a species relative to the other categories, where higher values indicate a stronger relationship.

Table 3-3 Summary of Predictive Diagnostic Coefficients in the Best-fit Generalised Linear Models for Four Faunal Groups at Roberts Bank

Predictor Variable Crustaceans Sea Stars Anemones Fishes Intercept (outside sea pen bed) -0.58* -0.43* -3.31* -1.20* Sea Pen Few to Patchy -0.12 -0.16 1.02* -0.16 Sea Pen Continuous to Dense 3.62* 3.56* 2.39* 2.97* Year - 2008 -0.04* -0.04* -0.10 -0.04 Year - 2011 -0.28* -0.35* -0.38* N/A Year - 2012 0.76* 0.65* 2.08* N/A Depth -0.50* -0.63* 3.72* -0.04* Notes: * indicates statistical significance (p< 0.05); N/A (not applicable) is assigned where coefficients presented in this table were not used in models for a particular faunal group. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 18 - November 2014

Crustacean presence data were best described by sea pen habitat, depth, and sampling year variables. Crustacean presence in areas with no sea pens showed a slightly negative association (-0.58; p<0.001), whereas a positive association (3.62; p<0.001) was observed with continuous dense sea pen habitat. No significant associations between crustacean presence and areas with few to patchy sea pens were noted (-0.12; p=0.24). Crustacean presence was also significantly, and slightly negatively, associated (-0.04; p<0.001) with depth while presence with sampling year was also significant, but varied from year to year in the direction of the relationship.

Sea star presence was best described by sea pen habitat and year variables. Similar to crustacean and fish results, areas with no sea pens and areas of continuously to densely distributed sea pens were significantly associated with sea star presence, showing negative (-0.43; p<0.001) and positive (3.56; p<0.001) coefficients, respectively. Again, no significant associations between sea star presence and areas with few to patchy sea pens were noted (-0.16; p=0.11). The association between sea star presence and sampling year was significant (p< 0.001) but not consistent, which may reflect variation among year in sampling.

Presence of the anemone group was best described by sea pen habitat, depth, and year variables. Anemone presence was significantly negatively correlated (-3.31; p<0.001) with areas of no sea pens, significantly positively associated (1.02; p< 0.001) with areas of few to patchy sea pens, and significantly positively associated (2.39; p= 0.02) in areas of continuous to dense sea pen distribution. Depth was significantly and slightly negatively associated (-0.10; p< 0.001) with anemone presence while year varied in the direction of the relationship. The ordinal regression model results for anemones suggests that there is a highly significant positive relationship between the distributional abundances of anemones and sea pens.

Fish presence was best described by sea pen habitat and depth variables. Similar to crustacean results, areas with no sea pens and areas of continuously to densely distributed sea pens were significantly associated with fish presence with negative (-1.20; p<0.001) and positive (2.96; p<0.001) coefficients, respectively. No significant associations between fish presence and areas with few to patchy sea pens were noted (-0.16; p=0.13). Fish presence was also slightly negatively, but significantly, associated with depth (-0.04; p<0.001). Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 19 - November 2014

Figure 3-4 Orange Sea Pen Distribution at Roberts Bank

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Figure 3-5 Slope Profiles along Orange Sea Pen Density Gradients

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Figure 3-6 Cumulative Distribution of Crustaceans from SIMS Surveys (2003, 2008, 2011) at Roberts Bank

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Figure 3-7 Cumulative Distribution of Sea Stars from SIMS Surveys (2003, 2008, 2011) at Roberts Bank

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Figure 3-8 Cumulative Distribution of Anemones, Sea Cucumbers, and Snails from SIMS Surveys (2003, 2008, 2011) at Roberts Bank

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Figure 3-9 Cumulative Distribution of Marine Fishes from SIMS Surveys (2003, 2008, 2011) at Roberts Bank Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 25 - November 2014

3.4.2 SCUBA Results

The intent of recording orange sea pen densities by diving rather than SIMS was to obtain better observational data due to sea pen retraction and expanding behaviour. Dive sites were located in the shallow, sub-tidal zone at depths of -3.5 to -5.0 m CD, and were selected to overlap with the depths of the densest portion of the sea pen bed. Data is presented in Table 3-4 (below).

3.4.2.1 Dive Site SP1

Dive Site SP1 was characterised by pronounced sand waves with less than 10% diatom cover and less than 10% shell debris that consisted of small shell fragments with some small and large half shells (butter clams (Saxidomus gigantea) and horse clams (Tresus sp.)). The only vegetation observed was drift eelgrass.

One orange sea pen was observed approximately two metres away from the anchor, but not in any quadrats. Tube-dwelling worms, likely species of the family Chaetopteridae (three-section tubeworms) and possibly the family Maldanidae (bamboo worms) were common, but not as abundant as observed at the other dive sites. Hermit crabs were also common. A few small flatfish (likely a species of sanddab) less than 5 cm long were observed.

3.4.2.2 Dive Site SP2

Dive Site SP2 was characterised by silty sand with fine wood debris and a 10 to 20% diatom cover. Scattered shell debris, with some areas of accumulated shell debris (25 to 50% cover), were observed and consisted of shell fragments as well as small and large half shells. Species that could be identified by the shells included cockles (Clinocardium sp.), butter clams, and likely horse clams. Drift eelgrass was also present.

As expected, the distribution of orange sea pens at this survey location was patchy. Densities ranged from 0 to 2/m2 with an average of 0.42 sea pens/m2 (n = 12). Most individuals were similar in size, approximately 30 to 40 cm tall, though some were withdrawn into their peduncle or main polyp and covered by a layer of silty sand, only identifiable by a slight depression in the substrate (similar to bivalve siphon depressions). Several smaller sea pens (approximately 10 to 15 cm tall) were observed.

Similar to SP1, tube-dwelling worms were abundant. Dungeness crabs were common, and were observed moving across the substrate or buried in the sand. Sunflower sea stars and small anemones attached to shell debris were also observed. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 26 - November 2014

3.4.2.3 Dive Site SP3

Dive Site SP3 was characterised by silty-sand with approximately 40% cover of diatoms. Shell debris (predominantly small shell fragments and half shells with some large half shells) was scattered throughout the area, and considered to be less than that observed at SP2 (no areas of accumulated shell debris observed). From what could be identified, the large half shells were butter clams and horse clams. The small half shells may be a species of Nutricola (dwarf-venus clam). Round/oval surface depressions in the seabed suggested bivalve presence; although siphons were not directly observed, some were observed retracting into the substrate and thought to be horse clams.

As expected, the distribution of orange sea pens at this site was continuous to dense. Densities ranged from 2 to 13/m2, with an average of 5.7/m2 (n = 12). Most sea pens appeared to be similar in size, approximately 30 to 40 cm tall. As observed at SP2, a large proportion of orange sea pens were withdrawn into their peduncle, some of which were covered by a layer of silty sand and only identifiable by a slight depression in the substrate.

Similar to the other two dive sites, tube-dwelling worms were abundant. Several Dungeness crabs were observed either moving across the substrate or buried in the sand. Other biota observed included hermit crabs, spiny pink stars, and small anemones attached to shell debris. Barnacles were also observed on the larger shell debris and were actively feeding. A sculpin (family Cottidae) and small shrimp were observed on one sea pen.

Table 3-4 Sea Pen Density Data for Dive Sites SP2 (sparse) and SP3 (dense) at Roberts Bank on October 19, 2011

Site Diver Depth (m); Time Quadrat # sea pens/m2

5 m; 10:30 1 0 2 1 3 2 Gina Lemieux 4 1

5 0

5 m; 10:41 6 0

Sparse

- 5 m; 10:30 1 0

SP2 SP2 2 0 3 1 Jamie Smith 4 0 5 0 5 m; 10:42 6 0 Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 27 - November 2014

Site Diver Depth (m); Time Quadrat # sea pens/m2 3.5 m; 11:36 1 7

2 2

3 2 Gina Lemieux 4 5

5 7

3.5m; 11:45 6 2

Dense

- 3.5 m; 11:34 1 13

SP3 SP3 2 6

3 4 Jamie Smith 4 5

5 7

3.5 m; 11:45 6 8 Note: No Sea Pens were Observed in Quadrats at Dive Site SP1 (absent) *Depths are reported relative to chart datum.

3.5 KEY FINDINGS

A discussion of the major results arising from the sea pen ecological study and data gaps is provided below.

3.5.1 Discussion of Key Findings

SIMS and SCUBA surveys of orange sea pen beds have enhanced the current understanding of the geographic extent and ecosystem interactions of orange sea pen at Roberts Bank. Results suggest that: i) the spatial extent of orange sea pens is greater than what has been previously documented; ii) aggregations of orange sea pens are not reproductively static, as previously understood; iii) there appears to be a general lack of natural orange sea pen predators; and iv) presence of associated fauna (crustaceans, sea stars, anemones, and fish) is more likely within areas of continuous to dense sea pens than areas where distribution is few to patchy or absent.

Prior to 2011, based on 2003 and 2008 SIMS surveys, the mapped extent of orange sea pen distribution was limited to a large field west of Westshore Terminals. SIMS results from 2011 indicated this area of continuous to densely distributed sea pens (~15 ha) was unchanged in spatial extent or density. In addition, the extension of survey boundaries northwest and into the existing dredge basin revealed the presence of a second dense aggregation (i.e., 7.6 ha in size) at the southern edge of Westshore Terminals, and scattered individuals in the existing dredge basin. It was not possible to distinguish whether the increase in spatial extent is a result of a growing population or simply reflective of expanded survey boundaries and sampling effort. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 28 - November 2014

Previous sea pen studies at Roberts Bank suggested this aggregation was reproductively inactive because of the absence of smaller sea pens (Hemmera 2009); however, juvenile sea pens (less than 15 cm height) were documented in 2011 (during SIMS and SCUBA surveys). The presence of multiple size (age) classes in 2011 indicates that these aggregations may represent a breeding population or, at least, offer conditions favourable for larval settlement, or both. These observations are consistent with literature that reports larval settlement can be patchy in space and highly episodic in time giving rise to discontinuous populations differing in age and size (Birkeland 1969, 1974).

Sea pen densities at dive sites SP2 (few to patchy) and SP3 (continuous to dense) increased between 2008 and 2011. At dive site SP2, mean orange sea pen density was higher in 2011 (0.42 sea pens/m2) than 2008 (0.08 sea pens/m2; n= 12). At dive site SP3 the increase was slighter, from a mean density of 4.3 sea pens/m2 in 2008 to a mean of 5.7 sea pens/m2 in 2011 (n = 12). Natural variation in the number of sea pens at each location between years is expected, considering dynamic oceanographic conditions, food availability, seasonality (July versus October), and reproductive strategies. Density differences may not be biological in origin but instead reflect sea pen retracting behaviour, which makes them imperceptible and underdetected.

The sea star-nudibranch-sea pen community is based on a known trophic network first described by Birkeland (1974). Known sea pen predators appear to be largely absent from orange sea pen aggregations at Roberts Bank, particularly species considered “sea pen specialists” (Birkeland 1974). This relief from predation pressure may explain, at least in part, how sea pens at Roberts Bank have been able to achieve such high numbers. Several leather and sunflower stars were observed within the patchy sea pen polygon, but no nudibranchs have been observed to date. Predation pressure has been shown to exert considerable influence on local sea pen abundance, with documented mortality rates as high as 97% from sea star and nudibranch feeding (Birkeland 1974). Predation patterns on sea pens at Roberts Bank may be different than those reported in Puget Sound studies, or perhaps predation does not play a major role in influencing sea pen abundance at this particular site.

The role of emergent fauna as physical habitat used by fish populations has been well studied for hard corals, particularly in tropical environments (Jones and Syms 1998, Auster 2005), but the association of fishes/macroinvertebrates with sea pens remains largely undefined. Correlations between abundance, or diversity, of organisms and their habitat have often been used as a measure of the importance of particular habitat features (Syms and Jones 2001) and the literature provides conflicting views of how closely coral reef fishes are associated with habitat variables (Jones and Syms 1998). Some authors have suggested corals are important for mediating fish distribution and abundance (Risk 1972), while others have demonstrated minimal associations of fishes with corals (Roberts and Ormond 1987, Syms and Jones 2001). Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 29 - November 2014

At Roberts Bank, significant trends between faunal presence and sea pen distribution were noted for all groups investigated (i.e., crustaceans, sea stars, anemones, and fish). Year and depth were included as variables to control for differences in the data collected (e.g., number of samples, locations etc.) and for differences in sea pen habitat types (“outside” the sea pen bed tends to be shallower), respectively. Results indicated that the likelihood of presence of each faunal group in continuous to dense sea pen habitat is significantly higher than in areas of either few to patchy or absent sea pen habitat (e.g., Photo 3; Appendix A); however, some variation in the direction and magnitude of the likelihood along the sea pen distribution gradient is likely (i.e., among the three sea pen habitat categories).

Although sea pen beds provide habitat used by a number of other species, there is not enough data to suggest a functional link between the habitat sea pens provide and demographic patterns of associated fish and macroinvertebrates. High densities of fishes in aggregations of sea pens do not necessarily indicate that sea pens provide a unique functional role, rather, they may simply have attributes similar to other important habitats. In other words, biological and geological habitats may be functionally equivalent for fish that favour structural relief. For example, Auster et al. (2005) demonstrated the false boarfish (Neocyttus helgae) used both fan-shaped corals and depressions in basalt pavement habitats as shelter or flow refuge. Fishes and structural fauna may co-occur in areas of high flows for enhanced prey delivery but have no direct association, as demonstrated by Tissot et al. (2006) with rockfishes (Sebastes sp.) in California and Koslow et al. (2000) with orange roughy (Hoplostethus atlanticus) in New Zealand.

While the level of orange sea pen importance in the demography of fish/invertebrate populations and communities remains to be established, a lack of studies demonstrating functional linkages between fish/invertebrate and sea pens does not imply that they do not play a role in mediating the distribution and abundance of associated species. Observations may be missing the time period when particular fishes/invertebrates (e.g., juveniles, spawners) use such habitats or that use of highly structured habitats is more spatially constrained or stochastic in nature (Auster 2007).

While the SIMS and SCUBA surveys have enhanced our understanding of the geographic extent and ecosystem interactions of orange sea pens at Roberts Bank, available information on this species can still be characterised as relatively data-poor. Similar to what Auster (2007) identified for deep-water corals, expanded observational studies of sea pens and non-sea pen features as shelter, sources of benthic prey, and sites with accelerated flows to enhance plankton prey delivery, are required to understand habitat linkages between orange sea pens, and fish/invertebrate populations and communities. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 30 - November 2014

3.5.2 Data Gaps and Limitations

Gaps in the survey data along the seaward edge of the slope, in deeper waters, exist due to: i) depth limit of the SIMS system, which was only capable of surveying to approximately 35 m depth; and ii) the presence of crab traps and commercial seine operations, which made navigation unsafe. Results from a Remotely Operated Vehicle (ROV) survey confirmed sea pen presence at depths up to -40 m CD (Hemmera 2014b), indicating that the full extent of sea pen distribution at Roberts Bank (including southeast to Point Roberts) remains undetermined.

The only literature available to compare to Roberts Bank sea pen density data is estimates from Puget Sound in the 1960s where Birkeland (1968, 1974) recorded densities of up to 22 sea pens/m2 in depths of -10 to -25 m. Sea pen density is notoriously difficult to estimate, as adult individuals tend to retract into their peduncle and can become completely unnoticeable. Caution is warranted in interpreting density results because it is difficult to accurately determine how many or what proportion of individuals are buried, leading to underestimation of abundance and density. Nevertheless, this study was able to meet the objectives of estimating the spatial distribution and density of sea pens at Roberts Bank, and documenting species associations.

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 31 - November 2014

4.0 LOCAL KNOWLEDGE STUDY

This section presents the methods and results of the orange sea pen local knowledge study.

4.1 STUDY METHODS

In an attempt to better understand the life history, spatial extent, and ecological importance of orange sea pens, a local ecological knowledge survey was developed and distributed to select individuals during January 2013. People with relevant direct knowledge, whether from a natural history or scientific/ecological analysis perspective, were identified based on desktop research and specific collegial referrals. Participants included academics, dive shop/tourism operators, fishermen, and aquarists.

Individuals were presented with a brief explanation of the proposed Project, the rationale and objectives of the questionnaire, and a request for participation in contributing their local knowledge. Approximately 85 individuals or organizations were contacted to participate in completing the questionnaire, of which 43 responses were received (51% response rate). Of the responses received, 31 individuals, henceforth referred to as ‘the participants’ contributed information specific to orange sea pens whether directly in the questionnaire or via email explanation, or by providing detailed data. Many of the remaining 12 responses did not contain information specific to orange sea pens but provided other useful information such as referrals to those who may be better-suited to provide relevant information.

The survey was structured so that questions pertaining to orange sea pens were presented in as logical and comprehensible a manner as possible. The four-page questionnaire was comprised of four sections with a total of 20 questions specific to orange sea pen life history, spatial extent, and role in supporting other marine species and ecological functions. The questionnaire was organised based on the following subjects:

 Geographical range and ecological significance;

 Abiotic drivers;

 Ascribed value; and

 Further information.

Appendix C provides a list of participants, list of questions and documented responses. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 32 - November 2014

4.2 RESULTS

4.2.1 Geographical Range and Ecological Significance

4.2.1.1 Species Distribution

Participants were asked to list and/or describe areas where they know orange sea pens exist. Responses corroborated what is reported in the literature and indicate that orange sea pens range from Alaska to California (Figure 4-1). Responses varied in terms of specificity, with some pointing to general coastal regions, while others pinpointed precise locations. Locally, in and around the Strait of Georgia, there were numerous accounts of orange sea pens in the Gulf Islands, Puget Sound, and Howe Sound (Figure 4-2).

4.2.1.2 Observed Densities

Participants were asked if they have observed dense aggregations of orange sea pens (defined as >four individuals/m2) and whether they would consider such aggregations to be a unique habitat feature. Of the 25 participants who directly observed orange sea pen aggregations along the Pacific Northwest coast, 16 individuals indicated that they had observed densities of >four individuals per m2 whereas 9 individuals had only observed densities of

Most participants who observed dense aggregations (>four individuals per m2) considered them to be a unique habitat feature of the area. Several responses indicated that orange sea pens can be a dominant feature on the seafloor, supporting the presence of other species that would not be there otherwise, thus contributing to overall species diversity; however, some participants did not consider sea pen aggregations particularly unique, citing that they occur at a number of locations under a variety of biotic and abiotic conditions

Several participants emphasised the difficulty in accurately estimating orange sea pen densities due to the species’ ability to bury into the surrounding sediment; consequently, examination of a site by divers tends to underestimate sea pen abundance and/or density.

4.2.1.3 Ecologically-Linked Species

Participants were asked whether they typically observe any fish or invertebrate species associated with orange sea pens. Most responses included nudibranchs and sea stars (known predators of orange sea pens). Other frequently observed species included, Dungeness, red rock (Cancer productus), and graceful rock (Metacarcinus gracilis) crabs, copper (Sebastes caurinus) and quillback (Sebastes maliger) rockfish, flatfish including starry flounder, and English (Parophrys vetulus) and C-O sole (Pleuronicthys coenosus), and geoduck clams. Participants were also asked to comment on whether they believe these species associations to be coincidental, or whether the species interact with one another. The most common response pointed to the role of orange sea pens as key prey items for certain species of nudibranch and sea stars. Multiple participants indicated that mobile species, such as crabs, rely on sea Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 33 - November 2014 pen beds for shelter/refuge while others deemed this relationship coincidental as they do not eat the pens or require their shelter. The geoduck–orange sea pen association; however, was considered to be coincidental as the species are thought to share the same habitat preferences but not interact.

Participants were then asked whether they have observed predators near orange sea pen colonies. Of the 24 participants that answered this question, 16 individuals indicated that they have directly observed nudibranchs preying on orange sea pens (particularly Tritonia diomedea, Tritonia festiva, and Armina californica). Many participants have also observed certain species of sea star (particularly Hippasteria spp.) feeding both on sea pens and on nudibranchs.

4.2.2 Abiotic Drivers

4.2.2.1 Substrate Type

Participants were asked to identify what type of substrate(s) orange sea pens prefer, based on their experience. There was consensus around sand being a preferred substrate, with all participants indicating they have observed orange sea pens in sand. The range of answers indicated that orange sea pens are commonly found in substrates ranging from silt-mud to sand-cobble with shell debris; however, several responses emphasised that orange sea pens are not found in silty habitats.

4.2.2.2 Flow

Participants were asked to describe the flow conditions in which they typically find orange sea pens. The majority of answers indicated sea pens are associated with moderate to high flow environments, with strong currents. One answer made the point that the largest colonies are always in areas of moderate to high current, and that the density drops off moving away from the current stream. Other answers indicated that orange sea pens can also be found in low-flow environments.

4.2.2.3 Depth

When asked what depth range they observe orange sea pens within, most participants answered between -5 to -30 m CD, but that they do occur deeper, commonly to -50 m. Several participants pointed out that the range identified may be limited to those depths accessible by SCUBA.

4.2.2.4 Unique Habitat Attributes

Participants were asked to describe any unique attributes of the surrounding environment where they observed orange sea pens, such as freshwater outflows. Approximately half of the participants noted that they have often found orange sea pens in estuarine habitats or other areas where freshwater sources are common. One participant made the point that while a number of known sea pen beds have nearby fresh water input, the work has not been done to establish that sites without freshwater input, but all the other substrate and flow characteristics, are less likely to have sea pens. The remaining half of participant answers indicated that they have not observed orange sea pens in the vicinity of freshwater outflows, or any other defining habitat attributes. This page is intentionally left blank. Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 34 - November 2014

Figure 4-1 Anecdotal Sea Pen Observations along the Pacific Coast

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 35 - November 2014

Figure 4-2 Anecdotal Sea Pen Observations in the Strait of Georgia

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 36 - November 2014

4.2.2.5 Significance of Abiotic Factors

Participants commented on whether they believe that any of the previously-mentioned abiotic factors dictate where orange sea pens choose to aggregate. Sandy substrate and the presence of moderate to strong currents were identified as the main factors that likely drive aggregation locations; however, several participants indicated that a combination of these abiotic factors dictates location choice.

One response indicated that, while sandy sediments are obviously the preferred sediment type, because individuals and low density populations occur in a variety of sediment types (ranging from silt to gravel pockets), this is not an absolute habitat-controlling factor. Another response suggested the patchy distribution of orange sea pens indicates that some factors might be favourable for targeting settlement, and that chemical cues from conspecifics (i.e., existing sea pens), along with substrate cues and possibly flow cues, might all contribute.

4.2.2.6 Additional Abiotic Factors

Participants commented on whether they were aware of, or have any hypotheses pertaining to, additional abiotic environmental characteristics that may control orange sea pen distribution and fitness. Possible hypotheses included salinity fluctuations, nutrient input (insofar as it affects plankton), light, and depth.

4.2.3 Ascribed Value

4.2.3.1 Ecosystem Functions

Participants commented on whether they have any hypotheses regarding important ecosystem functions (i.e., physical, chemical, and biological processes/attributes that contribute to the ecosystem) provided by colonies of orange sea pens. Many participants indicated that sea pens play a vital role as ecological engineers in that they turn over and oxygenate sediments through their burying behaviour, provide habitat structure and heterogeneity, and influence current flow over the substrate. The strong effects that sea pens might have on phytoplankton and sustaining food webs through nutrient cycling was also hypothesised by various participants.

4.2.3.2 Anthropogenic Value

Although most participants indicated that orange sea pens do not hold any specific role in their culture/lifestyle, every participant indicated that there is value in knowing that the species simply exists (existence value) and elaborated upon the concept of intrinsic value of biodiversity. Multiple participants indicated that, although the roles that orange sea pens play in the ecosystem are poorly understood, it does not make them unimportant. Several responses highlighted that these are fascinating and beautiful organisms that are a favorite amongst recreational divers.

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 37 - November 2014

4.2.3.3 Additional Comments

When participants were asked if there is any other information regarding P. gurneyi that they would like to contribute, 13 participants offered responses. Nine of the responses provided references to relevant contacts or journal articles. The remaining four responses, presented below, pertained to persistence of the species, highlighting both natural and anthropogenic causes for observed declines:

 “Beds don't seem to be a stable feature. They seem to persist for a couple decades and eventually get wiped out by predators; in the meantime new beds pop up (and many blink out while still juveniles) but some eventually take hold. I think many of the juvenile beds succumb to non-specialist predators like Hermissenda.” - Dr. Greg Jensen, University of Washington

 “Sea pens have been declining in Puget Sound. Some researchers who used to work on them there (e.g., near Golden Gardens State Park in Seattle) can’t find them in such abundance, if at all. I’m not sure if that is also true in the Strait of Georgia, but it does at least suggest that this species might be one of conservation concern.” - Dr. Chris Harley, University of British Columbia.

 “It is notable that Birkeland(1968:10) stated ‘In general, then, Ptilosarcus is never sparse.’ This statement was verified by Birkeland at a number of locations in Puget Sound. Thus, it is somewhat alarming that since approximately 1980, I have found the P. gurneyi populations in all of Birkeland’s original study areas to be sparse and only a relatively small fraction of their original density (Kyte 2001). However, the population that I recently observed on the west side of Ketron Island in South Puget Sound appeared to be nearly as dense as those studied by Birkeland. The only other location that may have dense P. gurneyi populations is Dash Point near Tacoma, Washington.” - Michael Kyte, Senior Marine Biologist.

 “It is really important to realise how little is known about the species, how important its biomass is, that it seems to be a key prey item to multiple species, that it is often in the shallows where it is particularly susceptible to the impacts of urbanization (chemical, physical disturbance), and – that research indicates it needs low flow areas whereby any development impacting flow is highly likely to have an impact (again particular susceptibility in the shallows).” - Jackie Hildering, Diver.

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 38 - November 2014

5.0 CLOSURE

Major authors and reviewers of this technical data report are listed below, along with their signatures.

Report prepared by: Hemmera Envirochem Inc.

Marina Winterbottom, Master of Marine Management Marine Biologist

Archipelago Marine Research

Pamela Thuringer, M.Sc., R.P.Bio. Marine Biologist

Report peer reviewed by: Hemmera Envirochem Inc.

Jamie Slogan, M.Sc., PhD (cand.), R.P.Bio. Senior Marine Biologist

Port Metro Vancouver Hemmera RBT2 – Orange Sea Pens - 39 - November 2014

6.0 REFERENCES

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Beaulieu, S. 2001. Life on glass houses: sponge stalk communities in the deep sea. Marine Biology 138:803–817.

Bell, S. S. 1985. Habitat complexity of polychaete tube-caps: influence of architecture on dynamics of a meioepibenthic assemblage. Journal of Marine Research 43:647–671.

Best, B. A. 1988. Passive suspension feeding in a sea pen: effects of ambient flow on volume flow rate and filtering efficiency. The Biological Bulletin 175:332–342.

Birkeland, C. 1968. Reciprocal interactions between a single prey species, Ptilosarcus gurneyi, and its complex of predators. Thesis, University of Washington, Seattle.

Birkeland, C. 1969. Consequences of differing reproductive and feeding strategies for the dynamics and structure of an association based on the single prey species, Ptilosarcus gurneyi (Gray). Disertation, University of Washington, Seattle.

Birkeland, C. 1974. Interactions between a sea pen and seven of its predators. Ecological Monographs 44:211–232.

Boutillier, J. A., E. Kenchington, and J. Rice. 2010. A review of the biological characteristics and ecological functions served by corals, sponges and hydrothermal vents, in the context of applying an ecosystem approach to fisheries. DFO Canadian Science Advisory Secretariat Research Document.

Buhl-Mortensen, L., and P. B. Mortensen. 2004. Crustaceans associated with the deep-water gorgonian corals Paragorgia arborea (L., 1758) and Primnoa resedaeformis (Gunn., 1763). Journal of Natural History 38:1233–1247.

Buhl‐Mortensen, L., A. Vanreusel, A. J. Gooday, L. A. Levin, I. G. Priede, P. Buhl‐Mortensen, H. Gheerardyn, N. J. King, and M. Raes. 2010. Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins. Marine Ecology 31:21–50.

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Burd, B. J., P. A. G. Barnes, C. A. Wright, and R. E. Thomson. 2008. A review of subtidal benthic habitats and invertebrate biota of the Strait of Georgia, British Columbia. Marine Environmental Research 66:S3–S38.

Burnham, K. P., and D. R. Anderson. 2002. Model selection and multimodel inference: a practical information-theoretic approach. Second Edition. Springer-Verlag, Berlin.

Chia, F. S., and B. J. Crawford. 1973. Some observations on gametogenesis, larval development and substratum selection of the sea pen Ptilosarcus gurneyi. Marine Biology 23:73–82.

Colpron, E., E. N. Edinger, and B. Neis. 2010. Mapping the distribution of deep-sea corals in the Northern Gulf of St. Lawrence using both scientific and local ecological knowledge. Canadian Science Advisory Secretariat (DFO).

Davis, N., and G. R. VanBlaricom. 1978. Spatial and temporal heterogeneity in a sand bottom epifaunal community of invertebrates in shallow water. Limnology and Oceanography 23:417–427.

DFO (Fisheries and Oceans Canada). 2007. Ecosystem overview: Pacific North Coast Integrated Management Area (PNCIMA). Edited by B.G. Lucas, S.M. Verrin, R. Brown for Canadian Technical Report of Fisheries and Aquatic Sciences.

DFO (Fisheries and Oceans Canada). 2011. Pacific Region cold-water coral and sponge conservation strategy (2010-2015). Fisheries and Oceans Canada.

DFO (Fisheries and Oceans Canada). 2012. Assessment of the impact of northern shrimp trawling on benthic habitats and communities in the estuary and northern Gulf of St. Lawrence. Science Advisory Report, Canadian Science Advisory Secretariat, Quebec Region.

Diaz, R., G. Cutter, and K. Able. 2003. The importance of physical and biogenic structure to juvenile fishes on the shallow inner continental shelf. Estuaries 26:12–20.

Dickinson, P. 1978. Conduction systems controlling expansion-contraction behaviour in the seapen Ptilosarcus gurneyi. Marine Behaviour and Physiology 5:163–183.

Edinger, E. N., V. E. Wareham, and R. L. Haedrich. 2007. Patterns of groundfish diversity and abundance in relation to deep-sea coral distributions in Newfoundland and Labrador waters. Bulletin of Marine Science 81:101–122.

Edwards, D., and C. Moore. 2008. Reproduction in the sea pen Pennatula phosphorea (Anthozoa: Pennatulacea) from the west coast of Scotland. Marine Biology 155:303–314.

Erhardt, H., and H. Moosleitner. 1998. Baensch Marine Atlas 2. Microcosm Ltd.

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FAO. 2008. Report of the technical consultation on international guidelines for the management of deep- sea fisheries in the high seas. Fisheries and Aquaculture Report, United Nations Food and Agriculture Organization, Rome, Italy.

Field, J. M., M. Yoklavich, J. Marignac, G. Caillet, R. Lea, and S. Bros. 2001. Deepwater habitat and fish resources associated with a marine reserve: implications for fisheries: Part II. Marine Ecological Reserves Research Program.

Fuller, S. D., F. J. Murillo Perez, V. Wareham, and E. Kenchington. 2008. Vulnerable marine ecosystems dominated by deep-water corals and sponges in the NAFO Convention area. Northwest Atlantic Fisheries Organization.

Gartner Lee. 1992. Environmental appraisal of proposed container terminal, Roberts Bank. Prepared for Vancouver Port Corporation.

Golder. 1996. Marine environmental review of a proposed grain terminal at Roberts Bank. Report for Vancouver Port Corporation.

Goodwin, C., and B. Pease. 1991. Geoduck, Panopea abrupta (Conrad, 1849), size, density, and quality as related to various environmental parameters in Puget Sound, Washington. Journal of Shellfish Research 10:65–77.

Gotshall, D., and L. L. Laurent. 1979. Pacific Coast subtidal marine invertebrates: A fishwatcher’s guide. Sea Challengers.

Harper, J. R., B. D. Bornhold, P. Thuringer, and D. McCullough. 1999. Application of underwater video imaging for seabed Engineering and habitat assessment. In proceedings of the 1999 Canadian Coastal Conference, Victoria, B.C.

Harper, J. R., B. Emmett, D. E. Howes, and D. McCullough. 1998a. Seabed imaging and mapping system - seabed classification of substrate, epiflora, and epifauna. In proceedings of the 1998 Canadian Hydrographic Conference, Victoria, B.C.

Harper, J. R., D. McCullough, B. Emmett, P. Thuringer, and A. Ledwon. 1998b. Seabed imaging and mapping system - pilot project results. Land Use Coordination Office, Victoria, B.C.

Heifetz, J. 2002. Coral in Alaska: distribution, abundance, and species associations. Hydrobiologia 471:19–28.

Hemmera. 2014a. Roberts Bank Terminal 2 technical report: Habitat suitability modelling study. Prepared for Port Metro Vancouver, Vancouver, B.C. in Port Metro Vancouver (PMV). 2015. Roberts Bank Terminal 2 Environmental impact statement: Volume 3. Environmental Assessment by Review Panel. Submitted to Canadian Environmental Assessment Agency..

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Hemmera. 2014b. Roberts Bank Terminal 2 technical data report: Marine invertebrates, marine fish & fish habitat - Marine benthic subtidal study. Prepared for Port Metro Vancouver, Vancouver, B.C. Available at: http://www.robertsbankterminal2.com/

Hixon, M. A., and B. N. Tissot. 2007. Comparison of trawled vs untrawled mud seafloor assemblages of fishes and macroinvertebrates at Coquille Bank, Oregon. Journal of Experimental Marine Biology and Ecology 344:23–34.

Hughes, D. 1998. Sea pens and burrowing megafauna (Volume III). An overview of dynamic and sensitivity characteristics for conservation and management of marine SACs. Scottish Association for Marine Science (UK Marine SACs Project).

Jones, G. P., and C. Syms. 1998. Disturbance, habitat structure and the ecology of fishes on coral reefs. Australian Journal of Ecology 23:287–297.

Kenchington, E., H. Link, V. Roy, P. Archambault, T. Siferd, M. Treble, and V. Wareham. 2011. Identification of Mega- and Macrobenthic Ecologically and Biologically Significant Areas (EBSAs) in the Hudson Bay Complex, the Western and Eastern Canadian Arctic. DFO Canadian Science Advisory Secretariat Research Document.

Koslow, J., G. Boehlert, J. Gordon, R. Haedrich, P. Lorance, and N. Parin. 2000. Continental slope and deep-sea fisheries: implications for a fragile ecosystem. ICES Journal of Marine Science: Journal du Conseil 57:548–557.

Kozloff, E. 1993. Seashore life of the northern Pacific Coast. University of Washington Press, Seattle.

Kyte, M. A. 2001. Vacant benthic habitats: where have all the sea pens gone? Pacific Estuarine Research Society.

Lamb, A., and B. P. Hanby. 2005. Marine life of the Pacific Northwest: a photographic encyclopedia of invertebrates, seaweeds and selected fishes. Harbour Publishing, Madeira Park, B.C.

Levin, L. A., and P. K. Dayton. 2009. Ecological theory and continental margins: where shallow meets deep. Trends in Ecology & Evolution 24:606–617.

Malecha, P. W., R. P. Stone, and J. Heifetz. 2005. Living substrate in Alaska: distribution, abundance, and species associations. Page 289 in. Volume 41. American Fisheries Society.

NOAA. 2010. Implementation of the Deep Sea Coral Research and Technology Program 2008-2009. Report to Congress.

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OSPAR. 2010. Background document for sea pen and burrowing megafauna communities. Biodiversity Series.

Pirtle, J. L. 2005. Habitat-based assessment of structure-forming megafaunal invertebrates and fishes on Cordell Bank, California. Thesis, Washington State University.

Risk, M. J. 1972. Fish diversity on a coral reef in the Virgin Islands. Smithsonian Institution.

Roberts, C., and R. Ormond. 1987. Habitat complexity and coral reef fish diversity and abundance on Red Sea fringing reefs. Marine Ecology Progress Series. Oldendorf 41:1–8.

Ryer, C. H., A. W. Stoner, M. L. Spencer, and A. A. Abookire. 2007. Presence of larger flatfish modifies habitat preference by Age-0 northern rock sole Lepidopsetta polyxystra. Marine Ecology Progress Series 342:227–238.

Ryer, C. H., A. W. Stoner, and R. H. Titgen. 2004. Behavioral Mechanisms Underlying the Refuge Value of Benthic Habitat Structure for Two Flatfishes with Differing Anti-Predator Strategies. Marine Ecology Progress Series 268:231 – 243.

Shimek, R. L. 2011. The life and death of sea pens. Reefkeeping Magazine 1–19. Available at http://reefkeeping.com/issues/2005-08/rs/feature/index.php

Soong, K. 2005. Reproduction and colony integration of the sea pen Virgularia juncea. Marine Biology 146:1103–1109.

Stoner, A. W., M. L. Spencer, and C. H. Ryer. 2007. Flatfish-habitat associations in Alaska nursery grounds: use of continuous video records for multi-scale spatial analysis. Journal of Sea Research 57:137–150.

Syms, C., and G. P. Jones. 2001. Soft corals exert no direct effects on coral reef fish assemblages. Oecologia 127:560–571.

Tissot, B. N., M. S. Love, K. York, and M. Amend. 2006. Benthic invertebrates that form habitat on deep banks off southern California, with special reference to deep sea coral. Fishery Bulletin 104:167– 181.

Triton. 2004. Deltaport Third Berth Project: Marine resources impact assessment. Triton Environmental Consultants Ltd., Richmond, B.C. Prepared for Vancouver Port Authority.

WoRMS. 2008. WoRMS taxon details: Ptilosarcus. World Register of Marine Species.

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7.0 STATEMENT OF LIMITATIONS

This report was prepared by Hemmera Envirochem Inc. ("Hemmera") and Archipelago Marine Research ("Archipelago"), based on fieldwork conducted by Hemmera and Archipelago, for the sole benefit and exclusive use of Port Metro Vancouver. The material in it reflects the authors' best judgment in light of the information available to them at the time of preparing this Report. Any use that a third party makes of this Report, or any reliance on or decision made based on it, is the responsibility of such third parties. Hemmera and Archipelago accept no responsibility for damages, if any, suffered by any third party as a result of decisions made or actions taken based on this Report.

Hemmera and Archipelago have performed the work as described above and made the findings and conclusions set out in this Report in a manner consistent with the level of care and skill normally exercised by members of the environmental science profession practicing under similar conditions at the time the work was performed.

This Report represents a reasonable review of the information available to Hemmera and Archipelago within the established Scope, work schedule and budgetary constraints. The conclusions and recommendations contained in this Report are based upon applicable legislation existing at the time the Report was drafted. Any changes in the legislation may alter the conclusions and/or recommendations contained in the Report. Regulatory implications discussed in this Report were based on the applicable legislation existing at the time this Report was written.

In preparing this Report, Hemmera and Archipelago have relied in good faith on information provided by others as noted in this Report, and have assumed that the information provided by those individuals is both factual and accurate. Hemmera and Archipelago accept no responsibility for any deficiency, misstatement or inaccuracy in this Report resulting from the information provided by those individuals.

APPENDIX A Photographs

Port Metro Vancouver APPENDIX A Hemmera RBT2 – Orange Sea Pens - 1 - November 2014

Photo 1: Sparse to Patchy Distribution of Sea Pens

Photo 2: Continuous to Dense Distribution of Sea Pens

Port Metro Vancouver APPENDIX A Hemmera RBT2 – Orange Sea Pens - 2 - November 2014

Photo 3: Dungeness Crab Hiding Under Orange Sea Pen

Photo 4: Retracted Sea Pens Indicated by Red Arrows

Port Metro Vancouver APPENDIX A Hemmera RBT2 – Orange Sea Pens - 3 - November 2014

Photo 5: Orange Sea Pens with Crab Trap

Photo 6: Juvenile Sea Pens Indicated by Red Arrows

APPENDIX B Statistical Summaries

Port Metro Vancouver APPENDIX B Hemmera RBT2 – Orange Sea Pens - 1 - November 2014

Predictive Diagnostic Predictor Variable Std. error z value p value Coefficient Crustaceans Intercept (outside sea pen bed) -0.58 0.07 -8.59 < 0.001 Sea Pen Few to Patchy -0.12 0.10 -1.18 0.24 Sea Pen Continuous to Dense 3.62 0.72 5.02 < 0.001 Year - 2008 -0.28 0.12 -2.35 0.02 Year - 2011 0.76 0.09 8.11 < 0.001 Year - 2012 -0.50 0.14 -3.63 < 0.001 Depth -0.04 0.00 -9.03 < 0.001 Sea Stars Intercept (outside sea pen bed) -0.43 0.06 -6.70 < 0.001 Sea Pen Few to Patchy -0.16 0.10 -1.60 0.11 Sea Pen Continuous to Dense 3.56 0.72 4.94 < 0.001 Year - 2008 -0.35 0.12 -2.98 0.00 Year - 2011 0.65 0.09 6.99 < 0.001 Year - 2012 -0.63 0.14 -4.57 0.00 Depth -0.04 0.00 -8.42 < 0.001 Anemones Intercept (outside sea pen bed) -3.31 0.16 -20.29 < 0.001 Sea Pen Few to Patchy 1.02 0.12 8.86 < 0.001 Sea Pen Continuous to Dense 2.39 1.04 2.30 0.02 Year - 2008 -0.38 0.20 -1.85 0.07 Year - 2011 2.08 0.16 12.92 < 0.001 Year - 2012 3.72 0.17 21.43 < 0.001 Depth -0.10 0.01 -14.78 < 0.001 Fishes Intercept (outside sea pen bed) -1.20 0.07 -18.36 < 0.001 Sea Pen Few to Patchy -0.16 0.11 -1.51 0.13 Sea Pen Continuous to Dense 2.97 0.76 3.92 < 0.001 Depth -0.04 0.00 -9.17 < 0.001

APPENDIX C Local Knowledge Survey Questionnaire and Results

ECOLOGICAL KNOWLEDGE QUESTIONNAIRE – ORANGE SEA PEN (PTILOSARCUS GURNEYI)

OBJECTIVE OF QUESTIONNAIRE:

On behalf of Port Metro Vancouver, Hemmera is seeking local knowledge pertaining to the marine fauna surrounding the proposed Roberts Bank Terminal 2 project in British Columbia. You have been identified as a participant based on a desktop study that identified individuals possessing relevant expertise, experience and/or knowledge in this particular field of study. Specifically, Hemmera is seeking your input in regard to known Orange Sea Pen (Ptilosarcus gurneyi) aggregations throughout the Pacific Northwest.

P. gurneyi are sessile colonial octocorals found in predominantly sub-tidal sandy bottom habitats (to over 100m) from southern California to Alaska (Gotshall and Laurent 1979). They are long lived species (to over 15 years) and they exhibit a spatially clumped pattern of recruitment(Birkeland 1974). Adult sea pens have the ability to retract into the sand at times becoming completely undetectable.

Although P. gurneyi has been documented sporadically along the west coast of Canada and the United States of America, limited current information exists regarding details on exactly where these invertebrates are found and their densities. Furthermore, studies describing the ecological importance of sea pen beds in near shore environments are lacking.

In an attempt to better understand the life history, spatial extent and ecological importance of P. gurneyi, we ask you to please take a moment to fill out the following survey to the best of your ability.

PART A. GEOGRAPHICAL RANGE AND ECOLOGICAL SIGNIFICANCE

1. Are you familiar with the orange sea pen, Ptilosarcus gurneyi? ▫ No (Please continue to Part D) ▫ Yes 2. Please describe the area along the Pacific Northwest coast in which you are familiar with P. gurneyi.

3. Please explain how you are familiar with P. gurneyi and the methods by which you have gathered information on them (e.g., direct dive surveys, studies of sea pens in lab setting, bycatch of sea pens in trawl gear, etc.).

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 2 - November 2014

4. Have you observed P. gurneyi directly?? ▫ No (Please continue to Part C) ▫ Yes, in aggregations of < 4 individuals per m2 ▫ Yes, in aggregations of > 4 individuals per m2 5. If you observed a dense aggregation of P. gurneyi (i.e. > 4 individuals per m2), would you consider this to be a unique habitat feature of the area? Please explain why and describe the density of the P. gurneyi aggregation that you observed (#’s per m2).

6. Did you observe juvenile fish or invertebrate species (e.g., rockfish, geoducks, shrimp, crabs) associated with P. gurneyi? Identified species can be associated with both small (<4 / m2) and large (>4 / m2) P. gurneyi aggregations. Please provide a description of associated species. ▫ No ▫ Yes i. ______ii. ______iii. ______7. Do you believe the above mentioned species associations to be coincidental? (i.e. both species simply prefer the same habitat)OR do you believe that these species have important interactions between one-another? (i.e. juvenile fish rely on sea pens as a refuge from predators)? Please explain.

8. Did you observe predators (i.e. nudibranchs and sea stars) near P. gurneyi colonies? Please describe interaction.

PART B. ABIOTIC (NON-LIVING) FACTORS

9. Were observed P. gurneyi found in sandy substrate? Please explain the observed substrate type.

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 3 - November 2014

10. Have you found P. gurneyi in high flow environments? Please explain.

11. At what depth range did you observe P. gurneyi?

12. Please describe any unique attributes of the surrounding environment where you observed P. gurneyi. Specifically, please comment on any freshwater outflows that may have been present in the vicinity.

13. Do you believe that any of the previously-mentioned abiotic (non-living) factors dictate where P. gurneyi choose to aggregate? Please explain.

14. Are you aware of, or have any hypotheses regarding additional abiotic (non-living) environmental characteristics that may control P. gurneyi distribution and fitness?

PART C. ASCRIBED VALUE

15. Do P. gurneyi (or sea pens in general) hold any specific traditional or cultural role for you? Please explain.

16. Is it important for you to know that P. gurneyi (or sea pens in general) simply exist, even if they do not play a role in your culture/lifestyle (i.e. existence value)? Please explain.

17. Are you aware of, or have any hypotheses regarding important ecosystem functions (physical, chemical, and biological processes/attributes that contribute to the ecosystem) attributable to aggregations of P. gurneyi?

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 4 - November 2014

PART D. FURTHER INFORMATION

18. Do you know of others who may be knowledgeable about P. gurneyi (or sea pens in general) along the Pacific Northwest coast? If yes, could you please provide their name and/or contact information? ▫ No ▫ Yes, please find name and/or contact information below: ______19. May we contact you via telephone to further discuss your knowledge regarding P. gurneyi? ▫ No thank you, I do not wish to be contacted further. ▫ Yes, you may contact me at the following number: ______20. Is there any other information regarding P. gurneyi that you would like to add? Please explain.

Thank you for taking the time to complete this survey on Ptilosarcus gurneyi.

Your input is much appreciated.

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 5 - November 2014

Table 1 Orange Sea Pen Local Ecological Knowledge Participant Information

# Name Organization Email 1 Dr. David Arsenault Bamfield Marine Science Center [email protected] 2 Dr. Charles Birkeland University of Hawaii [email protected] 3 Dr. Thomas Carefoot University of British Columbia emeritus [email protected] 4 Jim Cosgrove Royal BC Museum [email protected] 5 Dr. David Cowles Walla Walla University [email protected] 6 John DeBoeck Diver, Dive Industry Association of BC [email protected] 7 Dr. Barb Faggetter Ocean Ecology [email protected] 8 Donna Gibbs Vancouver Aquarium [email protected] 9 Dr. Chris Harley University of British Columbia [email protected] 10 Dr. Chris Harvey-Clark University of British Columbia [email protected] 11 Jackie Hildering Diver, Earthling Enterprises [email protected] 12 Dr. Greg Jensen University of Washington [email protected] 13 Chad King NOAA [email protected] 14 Michael Kyte Marine Biologist, The Talley Group [email protected] 15 Andy Lamb Marine Naturalist, Cedar Beach [email protected] 16 Steve Lonhart NOAA [email protected] 17 Patrick W. Malecha NOAA [email protected] 18 Peter Mieras Diver [email protected] 19 Roy Mulder Marine Life Sanctuary Society of BC [email protected] 20 Dr. James Murray California State University [email protected] 21 Melva Nikki Van Schyndel Echo Bay EcoVentures [email protected] 22 Bill Proctor (Referred by Salmon Coast Station) [email protected] 23 Dr. Ron Shimek Reef Stewardship Foundation [email protected] 24 Annette G. E. Smith Underwater Photographer [email protected] 25 Dane Stabel Diver [email protected] 26 Doug Swanston Diver [email protected] 27 Mike Tonnesen Diver [email protected] 28 Dr. VerenaTunnicliffe University of Victoria [email protected] 29 Dr. Gary Williams California Academy of Sciences [email protected] 30 Dr. Dennis Willows University of Washington [email protected] 31 Dr. Russell Wyeth St. Francis Xavier University [email protected]

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 6 - November 2014

Table 2 Associated Species Commonly Observed with Orange Sea Pens

Participants

Associated Species

Mieras

Russell Wyeth Russell Murray James Williams Gary Willows Dennis Ron Shimek Jensen Greg Cosgrove Jim DougSwanston Hildering Jackie Peter Smith Annette Birkeland Charles Kyte Michael Faggetter Barb VerenaTunnicliffe Malecha Pat Lonhart Steve Harvey Chris Clarke Nudibranchs Tritonia diomedea X X X X X X X X X X X

Tritonia festiva X X X X X X X X X

Armina californica X X X X X X X X X X X

Hermissenda X crassicornis Flabellina trophina X

Flabellina verrucosa X

Sea Stars Pynopodia X X X helianthoides Mediaster sp. X X X X X

Dermasterias X X imbricata Crossaster papposus X X X

Luidia foliolata X

Solaster sp. X X X

Pisaster brevispinus X

Hippasteria sp. X X X X

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 7 - November 2014

Participants

Associated Species

Mieras

Russell Wyeth Russell Murray James Williams Gary Willows Dennis Ron Shimek Jensen Greg Cosgrove Jim DougSwanston Hildering Jackie Peter Smith Annette Birkeland Charles Kyte Michael Faggetter Barb VerenaTunnicliffe Malecha Pat Lonhart Steve Harvey Chris Clarke Crabs Metacarcinus X X X X X X X X X magister Cancer productus X X

Metacarcinus gracilis X X

Pugettia productus X

Paralithodes X camtschaticus Pagurus armatus X

Rockfish Sebastes caurinus X X X X X

Sebastes maliger X X X

Sebastes nebulosus X

Flatfish Parophrys vetulus X X

Pleuronichthys X X coenosus Microstomus X pacificus Platichthys stellatus X X X

Citharichthys X sordidus Bivalves

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 8 - November 2014

Participants

Associated Species

Mieras

Russell Wyeth Russell Murray James Williams Gary Willows Dennis Ron Shimek Jensen Greg Cosgrove Jim DougSwanston Hildering Jackie Peter Smith Annette Birkeland Charles Kyte Michael Faggetter Barb VerenaTunnicliffe Malecha Pat Lonhart Steve Harvey Chris Clarke Panopea generosa X X X X X

Chlamys sp. X X

Tresus sp. X X

Other Cucumaria miniata X

Parastichopus X californicus Cymatogaster X aggregata Rossia pacifica X

Lunatia lewisii X X

Squalus suckleyi X

Virgularia sp. X

Hexagrammos X decagrammus Hydrolagus colliei X

Pandalus platyceros X

Crangon sp. X

Nassarius sp. X

Pachycerianthus sp. X

Table 3 Orange Sea Pen (Ptilosarcus gurneyi) Local Ecological Knowledge

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 9 - November 2014

Distribution Information

Recipient answers to Survey. Information consolidated from all sections of the survey plus emails. Responses are recorded verbatim.

Specific locations of P. gurneyi. This species is fairly common in most areas of nearshore sand/mud substrate from low intertidal to deep waters. In waters around Vancouver it is a common sand bottom species found in Howe sound and West Vancouver and in Indian Arm at such sites as Whytecliffe Park, Porteau Cove, Bowyer Island , and tends to favour areas with some current. Frequently Dr. Chris Harvey-Clarke, found in alrge patch aggregations but also singly or in small groups. The striped nudibranch is a common species that UBC parasitizes this species but I have not seen this species commonly in the Vancouver area, more so on the West coast of Vancouver island. Observations of concentrated aggregations in the regions of Prince Rupert Harbour, Chatham Sound, Browning Entrance, Dr. Barb Faggetter, 'Ocean Principe Channel, and Douglas Channel. Sporadic observations in many locations ranging from Dundas Island on the Ecology' North Coast to Klemtu on the Central Coast. One large bed is located in the Skeena River plume. Dr. Chris Harley, UBC Puget Sound, WA Associate Prof Puget Sound – From Whidby Is. South to Olympia, Dr. Ron Shimek, Prof, UW The American San Juan Islands – Waldron, Lopez, San Juan and Orcas. Friday Harbour Lab, BMSC

Vancouver Island – Saanich Inlet, Barkley Sound etc East end of Strait of Juan de Fuca/Whidbey & Fidalgo Islands. Dr. David Cowles, Walla I regularly observe them in Rosario Bay (only one small stream enters Rosario Bay and they are found about 100m from Walla Uni Prof the stream), which is an exposed location quite near Deception Pass. The highest concentration I have ever seen was off of Mayne Island in a moderate current/soft bottom (lots of shells) area. This area had the highest density of them I have ever seen. North Vancouver Island area also seems to have a fairly high Roy Mulder abundance. Passage Island reef (West Vancouver) seems to be a good area to find them. I may not be a biologist, but can provide some location data.  Central Puget Sound from Mukilteo to Tacoma including Blake Island, WA  South Puget Sound in the Nisqually Reach and around the western shore of Ketron Island, WA have dense aggregations Michael Kyte, Senior  The only other location that may have dense aggregations is Dash Point near Tacoma, Washington (location is near the Marine Biologist entrance to South Puget Sound).  North Puget Sound in the Cherry Point area  Roberts Bank, B.C. (through materials and information provided by Archipelago Marine Research, Ltd. in 2008) Washington State from Puget Sound, north to San Juan County, British Columbia from US boundary northwest outer coast Dr. Dennis Willows, Uni of to Cape Scott on Vancouver Island. Specifically, near McIntosh Rocks 5 km NW of Tofino. Washington Prof

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 10 - November 2014

Specific locations of P. gurneyi. Dr. Gary Williams, PhD, Gulf of Alaska, Puget Sound, Oregon to southern California. Curator Invert Zoology Puget Sound, San Juan Islands, Barkley Sound, Clayoquot Sound Dr. Russell Wyeth, St FX I have a number of tables and maps indicating both possible and known sites in the San Juan Islands and Puget Sound. asst. Prof My experience is mostly in the southern Vancouver Island area from Nanaimo to Sooke. There are several very large Jim Cosgrove, Royal BC fields in the waters surrounding Sidney and James Islands near Sidney. I have some specific area experience in Barkley Museum Sound (west Vancouver Island) and the Port Hardy area (north Vancouver Island). Pat Malecha, NOAA 2 dense aggregation around inside waters of Southeast Alaska near Juneau. Marine Ecology & Stock Assess Along Douglas Island between the city and Marmion Island…..about 6 miles long, large sea pen forests through this areaPoint Louisa Cove side large sea pen forestSunshine Cove – both sides of underwater reef – medium sized sea pen Annette Smith, Underwater forests. There are sea pen forests in areas around Coughlin Island, but I have not spent enough time underwater there to Photographer really discuss that area much. Dr. James Murray, I have dived a few times in sea pen beds off Burien WA, Bellingham WA, Lopez Sound WA, Shine, WA, and Tofino B.C. California State Uni Gulf Islands, Winchelsea islands, Johnstone Strait, Queen Charlotte Strait, Smith inlet, Kyuquot Sd, Barkley Sd, Nootka Sd, North coast channels (Hakai, Nalau, Seaforth, Millbank, Caamano, Principe, Otter, Beaver, Freeman) Haida Gwaii ( John deBoeck, Director of skidegate chan, Cumshewa Chan, Laskeek bay, Hoya Pass, Reef Island, Juan Perez Sd, Skincuttle Inl, Dolomite Dive Industry Asstn of BC narrows, Houston Stewart Channel, Anthony Island. I am familiar with their distribution within Monterey Bay (Monterey Bay Peninsula) and the northern Big Sur coast. Chad King, NOAA **INCLUDED A MAP AND EXCEL FILE REPRESENTING LOCATIONS. Plumper Island Group outside of Telegraph Cove, Hoya Head (Knight’s Inlet), Port Alice , “Seven Tree” near Browning Jackie Hildering, Diver Pass, Greenway Sound Areas of Saanich at 30 to 50m are dominated by whip corals and sea pens in appropriate sediments. Densities are not Dr. Verena Tunnicliffe, measured but we have the imagery that can support measurements. UVic Prof I have seen P. gurneyi in central California in the last couple of years, usually at a site near the entrance to Monterey Harbor. I possibly saw this species during dives on Santa Catalina Island, but it may have been at another site in southern Steve Lonhart, NOAA CA. I’d have to consult my notes if you really needed the info. Dr. Charles Birkeland, In Puget Sound near Seattle, mostly at Alki Point and Golden Gardens parks in Seattle. PhD, Prof (Uni Guam Marine Lab) West Coast Vancover Island (Bamfield) Dr. David Arsenault, BMSC

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 11 - November 2014

Specific locations of P. gurneyi. Barkley Sound Peter Mieras, Diver

Melva Nikki van Schyndel, The islands and inlets around the Broughton Archipelago, off the NE tip of Vancouver Island. Echo Bay. Naturalist in Echo Bay Bill Procter, Broughton Broughton Archipelago, especially Penphrase Pass with the highest density aggregation Archipelago Barkley Sound, Broughton Archipelago, Salish Sea, Howe Sound. Dane Stabel, Diver

Do see the sea pens up east side of Vancouver Island off Qualicum, and out around Hornby Island and inside Denman isl. Mike Tonnesen, Diver

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 12 - November 2014

Table 4 Orange Sea Pen (Ptilosarcus gurneyi) Local Ecological Knowledge

Recipient answers to Survey. Responses are recorded verbatim.

2 Please describe the area along the Pacific Northwest coast in which you are familiar with P. gurneyi. This species is fairly common in most areas of nearshore sand/mud substrate from low intertidal to deep waters. In waters around Vancouver it is a common sand bottom species found in Howe sound and West Vancouver and in Indian Arm at such sites as Whytecliffe Park, Porteau Cove, Bowyer Island , and tends to favour areas with some Dr. Chris Harvey-Clarke, current. Frequently found in alrge patch aggregations but also singly or in small groups. The striped nudibranch is a UBC common species that parasitizes this species but I have not seen this species commonly in the Vancouver area, more so on the West coast of Vancouver island. Observations of concentrated aggregations in the regions of Prince Rupert Harbour, Chatham Sound, Browning Dr. Barb Faggetter, 'Ocean Entrance, Principe Channel, and Douglas Channel. Sporadic observations in many locations ranging from Dundas Ecology' Island on the North Coast to Klemtu on the Central Coast. Dr. Chris Harley, UBC Puget Sound, WA Associate Prof Dr. David Cowles, Walla East end of Strait of Juan de Fuca/Whidbey & Fidalgo Islands Walla Uni Prof • Central Puget Sound from Mukilteo to Tacoma including Blake Island, WA • South Puget Sound in the Nisqually Reach and around Ketron Island, WA Michael Kyte, Senior • North Puget Sound in the Cherry Point area Marine Biologist • Roberts Bank, B.C. (through materials and information provided by Archipelago Marine Research, Ltd. in 2008) Washington State from Puget Sound, north to San Juan County, British Columbia from US boundary northwest Dr. Dennis Willows, Uni of outer coast to Cape Scott on Vancouver Island Washington Prof Dr. Gary Williams, PhD, Gulf of Alaska, Puget Sound, Oregon to southern California. Curator Invert Zoology Dr. Russell Wyeth, St FX Puget Sound, San Juan Islands, Barkley Sound, Clayoquot Sound asst. Prof My experience is mostly in the southern Vancouver Island area from Nanaimo to Sooke. I have some specific area Jim Cosgrove, Royal BC experience in Barkley Sound (west Vancouver Island) and the Port Hardy area (north Vancouver Island). Museum Pat Malecha, NOAA Inside waters of Southeast Alaska near Juneau. Marine Ecology & Stock Assess

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 13 - November 2014

Along Douglas Island between the city and Marmion Island…..about 6 miles long, large sea pen forests through this area Point Louisa Cove side large sea pen forest Annette Smith, Underwater Sunshine Cove – both sides of underwater reef – medium sized sea pen forests. Photographer There are sea pen forests in areas around Coughlin Island, but I have not spent enough time underwater there to really discuss that area much. I have 20 years of experience with a sea pen bed near Tacoma WA, and have also dived a few times in sea pen Dr. James Murray, beds off Burien WA, Bellingham WA, Lopez Sound WA, Shine, WA, and Tofino B.C. California State Uni Gulf Islands, Winchelsea islands, Johnstone Strait, Queen Charlotte Strait, Smith inlet, Kyuquot Sd, Barkley Sd, Nootka Sd, North coast channels (Hakai, Nalau, Seaforth, Millbank, Caamano, Principe, Otter, Beaver, Freeman) John deBoeck, Director of Haida Gwaii ( skidegate chan, Cumshewa Chan, Laskeek bay, Hoya Pass, Reef Island, Juan Perez Sd, Skincuttle Dive Industry Asstn of BC Inl, Dolomite narrows, Houston Stewart Channel, Anthony Island. I am not familiar with its distribution along the Pacific Northwest, but am familiar with their distribution within Chad King, NOAA Monterey Bay and the northern Big Sur coast. Plumper Island Group outside of Telegraph Cove, Hoya Head (Knight’s Inlet), Port Alice , “Seven Tree” near Jackie Hildering, Diver Browning Pass, Greenway Sound Dr. Verena Tunnicliffe, Saanich Inlet UVic Prof I have seen P. gurneyi in central California in the last couple of years, usually at a site near the entrance to Monterey Harbor. As I recall, I first learned about this species while studying in southern CA, and possibly saw it Steve Lonhart, NOAA during dives on Santa Catalina Island, but it may have been at another site in southern CA. I’d have to consult my notes if you really needed the info. Dr. Charles Birkeland, In Puget Sound near Seattle, mostly at Alki Point and Golden Gardens parks. PhD, Prof (Uni Guam Marine Lab) West Coast Vancover Island (Bamfield) Dr. David Arsenault, BMSC

As a diver regularly working as a diving marine biologist in British Columbia since 1981 between Southern British Doug Swanston, Diver Columbia, Bowie Seamount 150 Km West of Haida Gwaii and Prince Rupert. Seacology Barkley Sound Peter Mieras, Diver

I dive extensively Sidney to Victoria Harbour, Nanaimo north to Cape Lazo, Shearwater Spillar Inlet north including Mike Tonnesen, Diver Kitisoo Bay, Queen Charlottes, mainly greater Langara Island Melva Nikki van Schyndel, The islands and inlets around the Broughton Archipelago, off the NE tip of Vancouver Island. Naturalist in Echo Bay Bill Procter, Broughton Broughton Archipelago, especially Penphrase Pass with the highest density aggregation Archipelago Barkley Sound, Broughton Archipelago, Salish Sea, Howe Sound. Dane Stabel, Diver

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 14 - November 2014

The highest concentration I have ever seen was off of Mayne Island in a moderate current/soft bottom (lots of shells) area. This area had the highest density of them I have ever seen. North Vancouver Island area also seems Roy Mulder to have a fairly high abundance. Passage Island reef (West Vancouver) seems to be a good area to find them. I may not be a biologist, but can provide some location data. Puget Sound – From Whidby Is. South to Olympia, Dr. Ron Shimek, Prof, UW The American San Juan Islands – Waldron, Lopez, San Juan and Orcas. Friday Harbour Lab, BMSC Vancouver Island – Saanich Inlet, Barkley Sound etc Dr. Greg Jensen, Uni of Puget Sound / Salish Sea Washington Please explain how you are familiar with P. gurneyi and the methods by which you have gathered information on them (e.g., direct dive 3 surveys, studies of sea pens in lab setting, bycatch of sea pens in trawl gear, etc.). Dr. Chris Harvey-Clarke, Dive surveys UBC As an oceanographer and professional biologist, I am familiar with P. gurneyi from both an academic perspective and from field research in which I was carrying out marine benthic surveys. I have gathered information on them Dr. Barb Faggetter, 'Ocean primarily through the use of a georeferenced towed benthic video camera system which I have used to identify and Ecology' enumerate benthic organisms and classify benthic substrates. Dr. Chris Harley, UBC Invertebrate Zoology labs, talks by other researchers Associate Prof Professor of biology at Walla Walla University’s Rosario Beach Marine Lab near Anacortes, WA. Observed mainly Dr. David Cowles, Walla by diving + lab studies. Walla Uni Prof In the Puget Sound , I have had direct experience by conducting diving surveys. In some cases, these surveys were Michael Kyte, Senior for geoduck clams and in other cases for P. gurneyi. The surveys and observations have extended intermittently Marine Biologist from 1966 through 2011. Dive observations incidental to other work involving the seaslug predators of seapens, and otter trawl sampling and Dr. Dennis Willows, Uni of collecting in WA state. Washington Prof I am a specialist in pennatulacean systematics; geographical data is from our collections database at the California Dr. Gary Williams, PhD, Academy of Sciences, Department of Invertebrate Zoology and Geology. Curator Invert Zoology Dr. Russell Wyeth, St FX 200? SCUBA dives, ~500 hours of timelapse underwater video. asst. Prof I am a retired marine biologist with 20 years of professional employment at the Royal British Columbia Museum. All Jim Cosgrove, Royal BC my research was done by scuba diving however I also participated in studies that involved a variety of fishing and Museum dredging techniques. Pat Malecha, NOAA I have observed P. gurneyi while conducting scientific and sport dives. Marine Ecology & Stock Assess Annette Smith, Underwater Many years of diving. Collected Sea Pens for UC Davis (Dr. Dan Nurco) Photographer

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 15 - November 2014

I have explored the sea pen beds describe above as a means to collecting their predator Tritonia diomedea and related sea slugs, and have hand harvested as food for sea slugs I have collected. I have also spent much time Dr. James Murray, observing them in the lab, and do note them as bycatch in several trawls. I have done some chemical and California State Uni microscopic analysis of their tissues as well John deBoeck, Director of Direct diving- I ran a commercial dive live-aboard for recreational diver & photographers for over 20 years Dive Industry Asstn of BC I have observed them directly while SCUBA diving around Monterey Peninsula. I have also queried the MBARI VARS database for their ROV observations and CSUMB’s media records, which were either observed via ROV or Chad King, NOAA towed camera sled. I have included the map and both excel files representing the observations. Photo-documenting as a recreational diver. Jackie Hildering, Diver

Dr. Verena Tunnicliffe, Surveys of benthos by ROV UVic Prof I learned about the species as an undergraduate student at UCLA while taking field courses on Santa Catalina Island. I have also encountered the species while diving for my master’s thesis and doctoral dissertation research, Steve Lonhart, NOAA which focused on species within kelp forests, but I was often in sandy areas adjacent to kelp beds and saw the sea pen then. Dr. Charles Birkeland, Direct dive observations on their behavior and interactions with their predators. PhD, Prof (Uni Guam Marine Lab) Study of sea pen in laboratory setting Dr. David Arsenault, BMSC

Doug Swanston, Diver Direct Dive Surveys Seacology Observed while diving and filming Peter Mieras, Diver

Dive surveys for DFO and harvesting as a seafood harvester, commercial diver, as well as installations of docks. Mike Tonnesen, Diver

Melva Nikki van Schyndel, Personal study from books, scientific papers and articles from the Internet; bycatch from prawn fishing; diving Naturalist in Echo Bay Bill Procter, Broughton Visual sightings and observations at low tide, including counts of individuals Archipelago I encounter P. Gurneyi frequently while recreational and professional SCUBA diving, although I have not directly Dane Stabel, Diver surveyed or studied them. Dr. Ron Shimek, Prof, UW Direct diver surveys from 1973 through 1994. Laboratory studies of reproduction and embryology. Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of Dive surveys for other species (diving area since 1974) Washington

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 16 - November 2014

4 Have you observed P. gurneyi directly?? Dr. Chris Harvey-Clarke, Yes, in aggregations of both <4 and > 4 individuals per m2 UBC Dr. Barb Faggetter, 'Ocean Yes, in aggregations of both <4 and > 4 individuals per m2 Ecology' Dr. Chris Harley, UBC No Associate Prof Dr. David Cowles, Walla Yes, in aggregations of <4 individuals per m2 Walla Uni Prof Michael Kyte, Senior Yes, in aggregations of both <4 and > 4 individuals per m2 Marine Biologist Dr. Dennis Willows, Uni of Yes, in aggregations of <4 individuals per m2 Washington Prof Dr. Gary Williams, PhD, Yes, in aggregations of <4 individuals per m2 Curator Invert Zoology Dr. Russell Wyeth, St FX Yes, in aggregations of both <4 and > 4 individuals per m2 asst. Prof Jim Cosgrove, Royal BC Yes, in aggregations of both <4 and > 4 individuals per m2 Museum Pat Malecha, NOAA Yes, in aggregations of both <4 and > 4 individuals per m2 Marine Ecology & Stock Assess Annette Smith, Underwater Yes, in aggregations of both <4 and > 4 individuals per m2 Photographer Dr. James Murray, Yes, in aggregations of both <4 and > 4 individuals per m2 California State Uni Yes, usually less than 4 per m2 John deBoeck, Director of Can not recall much more than 4 per m2 Dive Industry Asstn of BC Yes, in aggregations of <4 individuals per m2 Chad King, NOAA

Yes, in aggregations of both <4 and > 4 individuals per m2 Jackie Hildering, Diver

Dr. Verena Tunnicliffe, Yes, in aggregations of both <4 and rarely > 4 individuals per m2 UVic Prof Yes, in aggregations of <4 individuals per m2 Steve Lonhart, NOAA

Dr. Charles Birkeland, Yes, in aggregations of both <4 and > 4 individuals per m2 PhD, Prof (Uni Guam Marine Lab)

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 17 - November 2014

No Dr. David Arsenault, BMSC

Doug Swanston, Diver Yes, in aggregations of both <4 and > 4 individuals per m2 Seacology Yes, in aggregations of both <4 and rarely > 4 individuals per m2 Peter Mieras, Diver

Yes, in aggregations of both <4 and > 4 individuals per m2 Mike Tonnesen, Diver

Melva Nikki van Schyndel, Yes, in aggregations of <4 individuals per m2 Naturalist in Echo Bay Bill Procter, Broughton Yes, in aggregations of <4 individuals per m2 Archipelago Yes, in aggregations of <4 individuals per m2, Although I have not directly observed high densities, I have boat tended for a researcher collecting Tritonia nudibranchs in Clayoquot Sound who reported high densities following Dane Stabel, Diver the dives. (See contact in section 18) Dr. Ron Shimek, Prof, UW Yes, in aggregations of both <4 and > 4 individuals per m2 Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of Yes, in aggregations of both <4 and > 4 individuals per m2 Washington If you observed a dense aggregation of P. gurneyi (i.e. > 4 individuals per m2), would you consider this to be a unique habitat feature of the 5 2 area? Please explain why and describe the density of the P. gurneyi aggregation that you observed (#’s per m ). Finding densities this high is not typical however I would estimate 20 percent of the observations I have made Dr. Chris Harvey-Clarke, would be of this high a density, usually in current swept areas. UBC Unfortunately, since P. gurneyi has never been a "species of interest" during any of our surveys, I do not have accurate density estimates for them. Typically, in a dense bed, I would observe 2 to 3 individuals per video frame. Since each video frame is approximately 0.2 m2, this would work out to be approximately 10 to 15 individuals per m2. Where P. gurneyi occurs at these densities, I definitely consider this to be a unique habitat feature of the area. Dr. Barb Faggetter, 'Ocean 2 In general, while I have observed P. gurneyi at low densities (e.g., much less than 1 individual per m ) commonly Ecology' throughout much of the North and Central Coast, I have only observed very dense aggregations at a small number of sites. Therefore, while no specific conservation issues have yet been flagged for P. gurneyi, I do comment on the presence of dense aggregations when I submit reports to clients based on the results of our benthic surveys. Yes, that seems like quite a dense aggregation. I typically find them at not more than 1-2/m2 though some can be Dr. David Cowles, Walla hiding below the sand. Walla Uni Prof I have observed “dense aggregations” of P. gurneyi where the measured population density was 10 to 30 individuals/m². These occurred along the eastern side of Puget Sound between Edmonds and Tacoma and in South Puget Sound on the western shore of Ketron Island. I would not consider that these relative dense populations were a “unique feature” because this “feature” occurred at a number of locations under a variety of biotic and abiotic Michael Kyte, Senior conditions. In addition, similar habitats (i.e., silty sand) with few or none P. gurneyi have been found. The most Marine Biologist recent (2011) relatively dense aggregation that I have observed was during geoduck stock assessment surveys in South Puget Sound. A notably dense aggregation was found along the western shore of Ketron Island. This population closely resembled the aggregations described by Birkeland (1974).

Port Metro Vancouver APPENDIX C Hemmera RBT2 – Orange Sea Pens - 18 - November 2014

I do not fully understand the question. Dense sea pen beds are patchily distributed, associated with a very Dr. Russell Wyeth, St FX particular set of flow and substrate features. So depending on how you define ‘area’ they may be unique (i.e. the asst. Prof only patch) or not. I have observed up to at least 20 per m2, probably higher. Yes, there are several very large fields of P. gurneyi in the waters surrounding Sidney and James Islands near Jim Cosgrove, Royal BC Sidney, B.C. These areas are mostly flat sand with moderate to strong currents. This area is commercially fished Museum for Cancer productus crab. I would estimate the density to vary from 4 per m2 to 8 per m2. I know of 2 aggregations of P. gurneyi near Juneau where density of the colonies are greater than or equal to 4 Pat Malecha, NOAA individuals per m2. In these areas, P. gurneyi is a dominant feature on the seafloor which could be considered Marine Ecology & Stock unique, Assess Where we have orange sea pens, they are fairly thick. We call them sea pen forests, as that is what they look like. However, viewing them depends on tides and currents, as they pull down into the ground when there isn’t much current running. The areas where orange sea pens reside, they are pretty much the only growing “plant type” sea Annette Smith, Underwater life there. White sea pens (aka sea whips) occupy areas deeper. So, normally, you will see a sea pen forest, pass Photographer through it, then a little deeper, the sea whips will start. I have lots of slides and photos of these areas. I have observed dense and sparse aggregations. The dense ones have had up to 10 per m2. In ideal habitat (flat, sandy,muddy bottom with adequate plankton to feed on), these densities are common, and the beds can persist for Dr. James Murray, decades. A sea pen bed is a “unique habitat” and supports the presence of many other associated species such as California State Uni sea slug predators, specific hermit crabs, flatfish, and crabs. When densities of sea pens are low, they do not have as an important effect on other species. I have not observed a dense aggregation. Chad King, NOAA

In my experience as a diver, these dense aggregations certainly appear to be rare and are associated with species like Triotonia festiva. They appear to have very specific habitat needs – low flow, a substrate high in sand and shell Jackie Hildering, Diver debris. The little clip I have made up at the following link gives a sense of this. http://themarinedetective.com/tag/diamondback-nudibranch/ “unique”? - not understanding the term. Areas of Saanich at 30 to 50m are dominated by whip corals and sea pens Dr. Verena Tunnicliffe, in appropriate sediments. Densities are not measured but we have the imagery that can support measurements. UVic Prof I have not seen dense clusters of P. gurneyi, but I would consider such an aggregation to be rare for the habitats I Steve Lonhart, NOAA have experience in, which includes the shallow subtidal (<30 m) in southern and central CA. Dr. Charles Birkeland, I do not understand what is meant by “unique habitat feature”. I did not estimate #’s per m2 because I had no way PhD, Prof (Uni Guam of knowing how many or what proportion of the population was withdrawn into the sand and out of sight. Marine Lab) Doug Swanston, Diver Yes, A unique assemblage of biota are associated with this habitat type. 4 and 20 P. gurneyi/m2 Seacology Approx 6 /m2 these are a few small areas of sandy bottom in medium current velocity . Peter Mieras, Diver

Bill Procter, Broughton Yes it would be unusual. Generally the aggregations are less dense and often you only see one on a reef. Archipelago

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I have seen this density commonly in virtually all of the areas I examined. Densities of adults are often quite a bit higher than seems so with a casual survey. The pens move up and down in the sediment burying themselves Dr. Ron Shimek, Prof, UW sometimes several times a day. Consequently, examination of a site by divers tends to underestimate the Friday Harbour Lab, BMSC abundances and densities. Adult abundances in many beds exceeds 10 per m2; Additionally, after settlement of etc the planktonic juveniles abundances can often be well in excess of 250 per m2 yes, because it influences overall species diversity- many seastars and nudibranchs that wouldn't be there if it was Dr. Greg Jensen, Uni of only a bare sand bottom.I don't have the data at hand for the surveys that measured density Washington Did you observe juvenile fish or invertebrate species (e.g., rockfish, geoducks, shrimp, crabs) associated with P. gurneyi? Identified species 6 can be associated with both small (<4 / m2) and large (>4 / m2) P. gurneyi aggregations. Please provide a description of associated species. Yes, i. Bivalve species especially geoduck, horse and other large bivalves and ii. sea whips where beds are found Dr. Chris Harvey-Clarke, in deeper water (25 m and deeper) are another common co aggregating species UBC Yes i. Copper rockfish (Sebastes caurinus) - adults ii. China rockfish (Sebastes nebulosus) - adults iii. Quillback rockfish (Sebastes maliger) - adults Dr. Barb Faggetter, 'Ocean iv. Geoduck clam (Panopea abrupta) Ecology' v. Scallop (Chlamys spp.) vi. Dungeness crab (Cancer magister) vii. Red sea cucumber (Cucumaria miniata) viii. California sea cucumber (Parastichopus californicus) Dr. David Cowles, Walla No Walla Uni Prof Yes • Juvenile Copper or Quillback rockfish vertically oriented to an individual P. gurneyi • Geoduck and gaper (Tresus sp.) are often associated with P. gurneyi beds. Michael Kyte, Senior • Cancroid crabs (Cancer sp. and Metacrinus sp.) Marine Biologist • The seastars Luidia foliata and Pycnopodia helianthoides • A variety of adult and juvenile flatfish including C-O Sole, Starry Flounder, and others • A variety of predators on P. gurneyi – see Question 8 Yes i. C.magister, ii. C.productus, Dr. Dennis Willows, Uni of iii. starry flounder, Washington Prof iv. English v. COsole, vi. seaslugs(Tritoniaand Armina) Dr. Gary Williams, PhD, Yes, Nudibranchs in the genus Tritonia. Curator Invert Zoology

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There are numerous associated species that I have observed. However, are you only interested in juveniles? Are you are trying to assess whether they are a nursery habitat? Proper identification of juvenile versus adults for the associated species is not something I have done. I expect many do not spend their entire lives in sea pen beds, but I do not have adequate information to delineate whether they are present as adult, juvenile or both. There are more than what I list below, however I would like clearer delineation of your criteria for including species in the list before I continue.: i. Tritonia diomedea ii. Armina californica iii. Virgularia sp. (in mixed pen and whip beds) iv. Pycnopodia helianthoides Dr. Russell Wyeth, St FX v. Solaster sp. asst. Prof vi. Mediaster sp. vii. Pisaster brevispinus viii. Zostera (in shallow areas) ix. Horse clams x. Geoduck xi. Ratfish xii. Sole xiii. Hermit crabs xiv. Cancer gracilis Yes i. A few rockfish (mostly S. carinus)_but more Kelp Greenlings Jim Cosgrove, Royal BC ii. Cancer magister crab_and a variety of smaller shrimp species Museum iii. Several very rare nudibranch species including Tritomia festiva, Tritonia diomedia and Armina californica Pat Malecha, NOAA Yes , Winter red king crab associations Marine Ecology & Stock Assess Yes, Dungeness Crab - Decorator crab really like being inside the sea pens and they even will decorate their shells with sea pens- hermit crabs, etc. One crab I almost never see among the sea pens is king crab. Not sure why, as they move through the areas….just normally they move deeper than the orange sea pens are, then come up Annette Smith, Underwater shallow.Wide variety of nudibranchs – in Sunshine Cove some of the largest tochuinas recorded have been found Photographer in there. The large nudibranchs generally do not mix. So….areas where diomedias reside with sea pens, you generally don’t see tochuinas (and vice versa)iii. Clams, scallops, mussels, etc.iv. I don’t see rockfish in their areas much, as the juvenile rockfish tend to be in the wallsv. _Sculpins, snailfish, gunnels, etc Yes i. flatfish like starry flounder ii. hermit crabs Dr. James Murray, iii. sea slugs Armina californica, Tritonia festiva, and Tritonia diomedea California State Uni iv. sea stars like Mediaster, Hipposterias v. Cancer magister I cannot recall specific species associations. Chad King, NOAA

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Yes i. Juvenile kelp crabs ii. Potentially juvenile copper rockfish Jackie Hildering, Diver iii. Sea slug predators mention in question 8 Yes?? “Associated” really should be a statistical test. my observations are idiosyncratic i. spot prawns Dr. Verena Tunnicliffe, ii. English and dover sole UVic Prof iii. dungeness crab Yes, as I recall, YOY rockfishes were in the vicinity, as well as sanddabs and possibly some scavenging snails, like Steve Lonhart, NOAA Nassarius Dr. Charles Birkeland, Yes There were some commensal associates such as tiny shrimp. I can send a closeup photo of one of the PhD, Prof (Uni Guam commensal shrimp if you wish. Marine Lab) o Yes .i. Tritonia diomedea Doug Swanston, Diver .ii. Tritonia festiva Seacology .iii. Armina californica YES .i. Striped and orange peel nudibranchs .ii. Dungeness and red rock crabs Peter Mieras, Diver .iii. Juv rock fish .iv. General sand bottom animals o Yes Mike Tonnesen, Diver i. sea stars, cukes, nudibranchs and other sandy bottom dwellers Melva Nikki van Schyndel, No Naturalist in Echo Bay Bill Procter, Broughton No Archipelago Yes i. There is an array of predators commonly found with P. gurneyi, (4-5 species of sea stars, and a like number of nudibranchs. See this link for more details: http://ronshimek.com/blog/?cat=218 ii. Additionally: geoducks present in beds where the density of adults is lower; they appear absent in dense Dr. Ron Shimek, Prof, UW beds.._Pachycerianthus common. Friday Harbour Lab, BMSC iii. Cancer gracilis and C. magister both are found in the beds, but not commonly. Crangon spp. shrimps, very etc common. Rossia pacifica present. Copper Rockfish, Quillback Rock fish present.; Squalus present, sometimes abundant.; Euspira lewisii, common. iv. Shiner perch common (Puget Sound areas). I haven't noticed any special associations with fish in our area. Geoducks often occur in the beds, but not at densities any higher than on open sand bottoms in the same general area; same goes for Pycnopodia, moon snails, Dr. Greg Jensen, Uni of and Pagurus armatus. Sea pen predators (and their predators) are much higher, so Hippasterias, Tritonia, Washington Mediaster, and Armina are usually abundant, and Crossaster is more common too.

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Do you believe the above mentioned species associations to be coincidental? (ie. both species simply prefer the same habitat) OR do you 7 believe that these species have important interactions between one-another? (I.e. juvenile fish rely on sea pens as a refuge from predators)? Please explain. Dr. Chris Harvey-Clarke, No data to support this but my impression is that the current conditions are what makes these species share habitat UBC In the case of sessile species (e.g., geoduck clam, red and California sea cucumbers), the species association with P. gurneyi is most likely coincidental, and probably reflects a preference for sandy habitats with moderate currents. In the case of motile species (e.g., rockfish and crab), the species association with P. gurneyi probably represents a beneficial species interaction. In flat, relatively featureless benthic habitats, I have often observed clusters of motile organisms (fish, shrimp, crab, squat lobsters) around a single tube worm, sea whip, or sea pen. Apparently, some organisms, such as prawn, are not usually associated with barren sediments, but appear to actively seek out habitats that are more complex [Schlining, K. L. 1999. The spot prawn (Pandalus platyceros Brandt 1851) resource Dr. Barb Faggetter, 'Ocean in Carmel submarine canyon, California: Aspects of fisheries and habitat associations. Moss Landing Marine Ecology' Laboratories. Stanislaus, California State University. M.Sc.: 54 pp.]. I suspect that there may be several possible explanations for these associations. Scavengers, such as crab and shrimp, may benefit from large particulate "food" which has been caught by a sea pen or tube worm, but rejected due to its large size. Some organisms, such as squat lobsters, may use sea pens, sea whips, or tube worms as "stakes" marking their territories. Small fish may use sea pens as refuges from predation, whereas larger fish may be benefiting from the current microeddies that form around the sea pen (reduces swimming effort or makes prey items easier to catch).  In the case of the juvenile rockfish associated with vertical expanded P. gurneyi, the individual fish were apparently using the P. gurneyi for shelter or refuge from predators. Juvenile and adult rockfish often seek out vertical objects with which to orient likely for refuge from predators and feeding. Thus, the association of juvenile rockfish with P. gurneyi is probably an opportunistic event.  Geoduck and gaper clams are often found in P. gurneyi beds likely because the sediment, water circulation, and Michael Kyte, Senior plankton characteristics are amenable to both species. This association is not obligatory because both clams and Marine Biologist sea pens are found without the other.  As with the geoduck and gaper clams, cancroid crabs, flatfish, and the seastars L. foliata and P. heliantoides co- occur with P. gurneyi likely because of biotic and abiotic habitat characteristics favorable and preferred by the species. Pycnopodia occasionally preys on P. gurneyi because the seastar is an opportunistic predator. Dr. Dennis Willows, Uni of Seaslugs eat the seapens. The other species may be coincidental. Washington Prof Dr. Gary Williams, PhD, These nudibranchs feed upon the sea pen Ptilosarcus gurneyi. Curator Invert Zoology The seastar-nudibranch-sea pen community is based on a known trophic network. Similarly, I think there are some Dr. Russell Wyeth, St FX bivalves and bivalve predators that co-exist. I would cautiously speculate that the geoducks and sea pens do not asst. Prof interact much, and thus the two trophic chains maybe somewhat independent and thus occur coincidentally. For the fish, shrimp and crabs I would say that it is a matter of food being abundant for all groups. There probably Jim Cosgrove, Royal BC is a prey/predator relationship betwenne the fish and small shrimps. The nudibranchs are predators on the sea Museum pens

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Pat Malecha, NOAA It is likely that the association is coincidental but also likely that the two species prefer similar substrates. Marine Ecology & Stock Assess Not so sure of the juvenile fish, However, I do know tochuinas really love to eat the sea pens. Diomedias also eat sea pens. They tend to not occupy the same places as the tochuinas. For example, you almost never see a diomedia in Sunshine Cove. There are lots of diomedias at Point Louisa yet you never see tochuina’s there. Annette Smith, Underwater Decorator crabs are often found hiding in the sea pens. They also use sea pens as part of their camouflage. Photographer Dungeness crab will often hide under the sea pens. Sometimes you will see snail fish hiding under seapens, however, only if there is a shell they can curl up in. The slugs depend on the pens as their exclusive food. The stars also seem to specialize on eating the pens nearly exclusively. The hermit crab species I see was cited in a crab book as being specifically associated with sea pen Dr. James Murray, beds. The flatfish and cancer crabs I think are mainly coincidental as they do not eat the pens or require their California State Uni shelter. There are also specific isopods that are symbionts that live inside sea pens. John deBoeck, Director of Juvenile fish DO rely on sea pens for refuge & shelter Dive Industry Asstn of BC I do not have enough information to answer this. Chad King, NOAA

Not coincidental for sea slugs – key prey item. Jackie Hildering, Diver

It is likely that sea pens provide the bottom roughness that some species seek – we have not done the tests Dr. Verena Tunnicliffe, necessary. UVic Prof In some cases, the sea pen provides habitat to mobile species, and especially small fishes. I am not sure if this results in net positive production for those species that opportunistically use the sea pen vs. a simple redistribution Steve Lonhart, NOAA of the species that are already there. Dr. Charles Birkeland, Although Ptilosarcus is preyed upon by at least seven species of predators (four asteroids and 3 nudibranchs), I PhD, Prof (Uni Guam suspect that only Hippasteria, Armina and Tritonia are associates. Marine Lab) Doug Swanston, Diver Interspeceis interactions including preditor prey Seacology The striped and orange peel nudibranchs feed on the orange sea pen Peter Mieras, Diver

I think coincidental; do not see fish interacting with sea pens Mike Tonnesen, Diver

Bill Procter, Broughton Unsure Archipelago Obviously, the predatory array found with the pens is not coincidental. Probably none of the other associations are Dr. Ron Shimek, Prof, UW incidental either. The bioturbation of the habitat by the pens severely disturbs the substrate, and this seems to Friday Harbour Lab, BMSC elimate most other sediment dwelling species such as clams after a period of time. There are probably other etc important interactions, but this is the big one. Geoducks, Pagurus armatus, Pycnopodia and moon snails appear to be coincidental; others mentioned above are Dr. Greg Jensen, Uni of either eating the pens or (in the case of Crossaster) eating the nudies. Washington

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8 Did you observe predators (i.e. nudibranchs and sea stars) near P. gurneyi colonies? Please describe interaction. Dr. Chris Harvey-Clarke, See above comment 2 re striped nudibranch UBC Yes, I observed predators near P. gurneyi colonies. While most sea star species generally occurred both near P. gurneyi colonies and at locations where there were no P. gurneyi, I definitely observed concentrations of sea stars in and around some of the colonies. Specific species of sea stars which I have seen in high abundance around P. gurneyi colonies are rose stars (Crossaster papposus), painted stars (Orthasterias koehleri), sunflower stars Dr. Barb Faggetter, 'Ocean (Pycnopodia helianthoides), long ray stars (Stylasteria forreri), false ochre stars (Evasterias troschelli), ochre stars Ecology' (Pisaster ochraceus), and short-spined stars (Pisaster brevispinus). I have also observed white-lined nudibranchs (Dirona albolineata) feeding on P. gurneyi. In this situation, the nudibranchs were not found outside the P. gurneyi colony at the study site. I have not directly observed predation but am well aware of several local predators and have seen the results of Dr. David Cowles, Walla predation (stripped pens) Walla Uni Prof During my work with Charles Birkeland (1968, 1974) and in later years and other locations, I have frequently encountered species that are documented predators of P. gurneyi. On occasion, these species, especially the sea stars, have been observed in the absence of P. gurneyi. Where P. gurneyi were present, the predators were Michael Kyte, Senior observed feeding on P. gurneyi, foraging between individuals P. gurneyi, and mating or laying egg masses. I have Marine Biologist observed nudibranchs, especially Armina, and seastars, particulary Hippasteria and Solaster sp., actively feeding on P. gurneyi. Dr. Dennis Willows, Uni of Yes, nudibranchs, hunt and eat seapens Washington Prof Dr. Gary Williams, PhD, Yes, see 6 & 7 above Curator Invert Zoology Many times. Tritonia grazes on Ptilosarcus, leaving the bulk of the colony unharmed. Packs of Armina consume Dr. Russell Wyeth, St FX entire Ptilosarcus. My colleagues and I have observed Hippasterias wipe out an entire sea pen bed, as described asst. Prof by Birkeland. Yes, see above answer regarding nudibranchs. No sea stars that I recorded. I have photos of some of the Jim Cosgrove, Royal BC nudibranchs feeding on P. gurneyi. Museum Pat Malecha, NOAA I have observed the nudibranch Tritonia festiva. Marine Ecology & Stock Assess See above. Tochuinas and diomedias enjoy dining on sea pens. I see lots of baby basket stars but they generally Annette Smith, Underwater are not found around the orange sea pens. They tend to use the larger white sea whips for protection. Our orange Photographer sea pens are fairly shallow, while the sea whips live deeper. Yes, the 3 nudibranchs above eat the sea pens as their exclusive food source. See above for sea stars predation Dr. James Murray, too. One sea pen bed I observed for 20 years was recently eliminated, and this coincided with an increase in California State Uni Hipposterias predation. John deBoeck, Director of Yes- toquerina tetraquatra eat them Dive Industry Asstn of BC

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I do not have enough information to answer this. Chad King, NOAA

I have photographed Tochuina tetraquetra and Triotonia festiva feeding on orange sea pens. I have observed Jackie Hildering, Diver Armina californica doin 8g so. Painted sea stars, leather star rose star, morning sun star. Dr. Verena Tunnicliffe, insufficient observations UVic Prof I have seen Dendronotus iris near P. gurneyi. Steve Lonhart, NOAA

Dr. Charles Birkeland, You cited my paper “Birkeland 1974”, and that describes all I can remember. PhD, Prof (Uni Guam Marine Lab) Doug Swanston, Diver Yes Seacology Striped and orange peel nudibranchs feeding on the orange sea pen Peter Mieras, Diver

I do see sea stars and nudibranchs but do not recall interactions Mike Tonnesen, Diver

Melva Nikki van Schyndel, Yes, I saw a sunflower star wrapping itself on it and so I assume it was attempting to eat it in their upright form. Naturalist in Echo Bay Bill Procter, Broughton No Archipelago Of course, again see the link: http://ronshimek.com/blog/?cat=218 Asteroids: Hippasteria spinosa, Crossaster papposus, Dermasterias imbricata, Mediaster aequalis = all observed Dr. Ron Shimek, Prof, UW eating pens. Friday Harbour Lab, BMSC Nudibranchs: Armina californica, Tritonia festiva, Tritonia diomedea, Tritonia tetraquetra, all observed eating adults. etc Hermissenda crassicornis, Flabellina trophina, Flabellina verrucosa – commonly in the beds, presumably eating juvenile sea pens. Dr. Greg Jensen, Uni of Many times- Hippasterias and Armina feeding on pens. Have rarely caught Tritonia or Mediaster in the act. Washington

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PART B. ABIOTIC FACTORS 9 Were observed P. gurneyi found in sandy substrate? Please explain the observed substrate type. Dr. Chris Harvey-Clarke, Yes, shell sand and mud ss UBC I have observed P. gurneyi in substrates ranging from silt-mud to sand-cobble. Most frequently, large P. gurneyi Dr. Barb Faggetter, 'Ocean colonies have occurred in silt-mud to sand substrates. Ecology' Dr. David Cowles, Walla Yes, primarily sand Walla Uni Prof P. gurneyi that I have observed have most often been associated with sandy sediments with varying degrees of Michael Kyte, Senior finer sediments (i.e., silt). I have also observed single P. gurneyi in gravel or rocky habitats where the P. gurneyi Marine Biologist occupied a pocket of sandy or silty gravel. Dr. Dennis Willows, Uni of Sandy and muddy substrates, yes Washington Prof Dr. Gary Williams, PhD, Yes, sandy or other soft bottom sediments. Curator Invert Zoology Sea pens seem to prefer shallow shelving sand-mud mixed substrates. Shell content seems to vary in my Dr. Russell Wyeth, St FX experience. Steep shelving areas also occasionally (but less frequently) support high density beds. Flat and asst. Prof muddy substrates tend to have more Virgularia (if any pennatulaceans are present at all). Jim Cosgrove, Royal BC Yes, the majority have been found either in sand or in mixed sand/mud substrate. Museum Pat Malecha, NOAA The P. gurneyi aggregations are in areas with sandy substrates with little vertical relief. Marine Ecology & Stock Assess In most cases our waters have a silty bottom. Sea pens live in the silty areas. They do not live on rock walls, and I Annette Smith, Underwater have not seen them in the slides on walls. Photographer Dr. James Murray, Yes, sandy/muddy substrate with a substantial diatom mat. California State Uni John deBoeck, Director of Sandy and/or sand/shell and/or sand/shell/pebble Dive Industry Asstn of BC I directly observed them in sandy substrate Chad King, NOAA

Yes –dense aggregations appear to be if there is high shell debris in sand. Jackie Hildering, Diver

Dr. Verena Tunnicliffe, silt to fine sand UVic Prof Yes, usually fairly fine sediment. Steve Lonhart, NOAA

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Dr. Charles Birkeland, Yes. Sandy. PhD, Prof (Uni Guam Marine Lab) Doug Swanston, Diver Yes Seacology Sandy bottom Peter Mieras, Diver

Sandy bottoms and slightly cobbled, gravel Mike Tonnesen, Diver

Melva Nikki van Schyndel, Yes it was fine sandy. At points more muddish in texture Naturalist in Echo Bay Bill Procter, Broughton They were observed on a flat, rocky reef with shell pieces, at depth of about 4 feet at low tide. Archipelago I have seen them in sandy and coarser shell substrates. Dane Stabel, Diver

Dr. Ron Shimek, Prof, UW I have found them in sand, shell fragment gravel, sandy silt. They don’t like really silty habitats. Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of Almost always in fairly clean sand (no silt or mud). Occasionally in areas with gravel mixed in. Washington 10 Have you found P. gurneyi in high flow environments? Please explain. Dr. Chris Harvey-Clarke, defintley – moderate currents for the most part- extreme current environments not so much UBC Yes, I have found P. gurneyi in high flow environments. As an example, one P. gurneyi colony that we have surveyed was located at the end of a point which had high tide and wave activity. Another P. gurneyi colony was Dr. Barb Faggetter, 'Ocean located in a shallow region of Prince Rupert harbour, where the high tidal range generates very strong currents in Ecology' the shallows. Yes, in moderately high flow. I regularly observe them in Rosario Bay, which is an exposed location quite near Dr. David Cowles, Walla Deception Pass. Currents in the bay itself don’t often exceed 1 knot but are seldom completely still. Walla Uni Prof I have occasionally found P. gurneyi in relatively high flow conditions. The most recent of these was in South Puget Sound where P. gurneyi was found in relatively coarse well-sorted sand that was heavily rippled by strong tidal Michael Kyte, Senior currents. In addition, the photographic materials provided to me by Archipelago Marine Research, Ltd. In 2008 Marine Biologist clearly showed P. gurneyi in an area subject to strong currents. The relative current strength was evident from the inclined position of the individual P. gurneyi, sand ripples, and the behavior of the scuba divers. Dr. Dennis Willows, Uni of Yes near McIntosh Rocks 5 km NW of Tofino Washington Prof Dr. Gary Williams, PhD, No Curator Invert Zoology

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This is hard to answer. Certainly, I have experienced a few sites where flow approach dangerous speeds for SCUBA. However, I have not observed them around Race Rocks, which has very high flow. But then, diving in Dr. Russell Wyeth, St FX higher flow environments is more restricted to rocky substrates where the topography provides local refuges for asst. Prof SCUBA divers. The largest colonies are always in areas of moderate to high current. The density drops off the farther you are out Jim Cosgrove, Royal BC of the current stream. Places of low current such as Saanich Inlet have some P. gurneyi but the colonies are very Museum isolated and often number less than 10 individuals. Pat Malecha, NOAA No, the areas are affected by tidal exchanges. Marine Ecology & Stock Assess Define high flow. Yes, they tend to like areas of current but not rushing like a river. They will generally be hidden Annette Smith, Underwater when the waters are calm and rise up when there is water movement. Note…I said generally, not usually or Photographer always! Dr. James Murray, Yes, they require high flow, ranging from 20-100 cm/s. California State Uni YES- most higher density colonies are in waters which move at > 1 m/sec max flow- and up to 6 m/sec max flow John deBoeck, Director of They seem to ‘thrive’ in waters which move at 1 to 3 m/sec (max flow speed on largest tides) Dive Industry Asstn of BC Not personally Chad King, NOAA

No Jackie Hildering, Diver

Dr. Verena Tunnicliffe, No UVic Prof I have not noted that, but then I tend to avoid high flow environments while diving. Steve Lonhart, NOAA

Dr. Charles Birkeland, Yes, but also in low-flow environments. PhD, Prof (Uni Guam Marine Lab) Doug Swanston, Diver Yes Seacology Medium velocity areas Peter Mieras, Diver

do not recall, but work a lot in these areas and it does not stand out Mike Tonnesen, Diver

Medium flow. But mostly slow/calm flow environments. No rapids but strong currents at times moving through Melva Nikki van Schyndel, channel gap (+100ft) Naturalist in Echo Bay Bill Procter, Broughton No Archipelago Dr. Ron Shimek, Prof, UW Occasionally, but only in areas where they have been protected from major current flow (i. e. in the lee of large Friday Harbour Lab, BMSC rocks). etc Yes, but I don't have measurements of flow speed. Strong enough that it is difficult to swim against, so probably Dr. Greg Jensen, Uni of about a knot. In some of the higher flow places the bottom tends to be more gravelly and I see juveniles but not Washington adults.

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11 At what depth range did you observe P. gurneyi? Dr. Chris Harvey-Clarke, 6m to 45 m UBC I have observed P. gurneyi at depths ranging from 5 m to 65 m. The most common depth range for P. gurneyi that Dr. Barb Faggetter, 'Ocean we have surveyed has been from 10 m to 30 m. Ecology' Dr. David Cowles, Walla Approximately 8-10 m Walla Uni Prof Michael Kyte, Senior I have observed P. gurneyi from the intertidal zone down to over 50 meters MLLW. Marine Biologist 15m to 3m, along shorelines usually, but less often down to 35 . Could reflect my diving depth habits and Dr. Dennis Willows, Uni of constraints. Washington Prof Dr. Gary Williams, PhD, Approximately 25 ft. – 225 ft. Curator Invert Zoology Dr. Russell Wyeth, St FX 2m – 30m by SCUBA. I have alse seen a few trawled from deeper, but have not spent much time doing that. asst. Prof Jim Cosgrove, Royal BC Normal scuba diving depths from 1 meter to 30 meters. Museum Pat Malecha, NOAA 10-30 m Marine Ecology & Stock Assess Usually between 30 and 80 feet. Most of them are in the 35 to 60 foot range. I have never seen them at the 100 Annette Smith, Underwater foot level….but then, I am usually diving walls at that depth and these guys don’t hang out on walls!! I never see Photographer sea pens near walls, even in the slide areas of a wall. Dr. James Murray, Usually not found shallower than 5 m MLLW, and not often deeper than 30 m. California State Uni John deBoeck, Director of 7 m to 45 m Dive Industry Asstn of BC 40-80 feet. Further depth ranges can be found within the attached files. Chad King, NOAA

From about 15 feet to about 80 feet (limited only by my diving depth I believe – likely deeper). Seem to do Jackie Hildering, Diver particularly well in the shallow – forming “fields”. Dr. Verena Tunnicliffe, 30 to about 60 m but can get better range if necessary UVic Prof From 6-30 m deep. Steve Lonhart, NOAA

Dr. Charles Birkeland, As deep as I could dive, i.e., about 150 ft. PhD, Prof (Uni Guam Marine Lab)

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Doug Swanston, Diver to 30 m RTCD Seacology From 5meters down to 20 meters Peter Mieras, Diver

generally 10 to 40 feet Mike Tonnesen, Diver

Melva Nikki van Schyndel, approx 30ft and at low tide some areas are approx 10 ft Naturalist in Echo Bay Bill Procter, Broughton 3-4 feet depth at low tide but they are often much deeper. Archipelago 20-100ft Dane Stabel, Diver

Dr. Ron Shimek, Prof, UW US datum: -0.3m to -50m; trawled from – 200 m. Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of Generally from 20-80 ft., but that's the depth for most of my dives. Washington Please describe any unique attributes of the surrounding environment where you observed P. gurneyi. Specifically, please comment on any 12 freshwater outflows that may have been present in the vicinity. Most areas I dive are away from fw sources due to sediment and diminished visibility so this would be hard to Dr. Chris Harvey-Clarke, determine UBC The largest aggregations of P. gurneyi that I have observed have been in environments with low slope, low rugosity, and moderate to high currents. Several of the largest aggregations have been associated with some amount of Dr. Barb Faggetter, 'Ocean fresh water input - one large bed is located in the Skeena River plume, another is located near a small creek, and a Ecology' third is close to a waterfall exiting from a freshwater lake. Only one small stream enters Rosario Bay. The P. gurneyi are first found about 100 m from the stream. The Skagit Dr. David Cowles, Walla River empties into the ocean at the other end of Deception Pass so much freshwater enters the ocean nearby. Walla Uni Prof Salinity near the sea pen colony in the summer is about 29 ppt. I cannot think of any unique attributes of the habitats in which I have found P. gurneyi. All the habitats have been estuarine by definition. In particular, South Puget Sound receives the outflow of a number of streams including the Michael Kyte, Senior relatively large Nisqually River on whose delta I have seen P. gurneyi. The most characteristic P. gurneyi habitat Marine Biologist feature is the silty sand in which they are most commonly found. Dr. Dennis Willows, Uni of Often there are freshwater outlets nearby. Washington Prof Dr. Gary Williams, PhD, N/A Curator Invert Zoology I have spent considerable energy attempting to identify candidate sites for sea pen beds. The factors I came to focus on were: some flow, sand or mud substrate, and shelving substrate (i.e. not flat enclosed bays, nor steep Dr. Russell Wyeth, St FX rocky drop-offs). A number of known sea pen beds have nearby fresh water input. However, I do not believe asst. Prof anyone has done the critical comparison to establish that sites without freshwater input but all the other substrate and flow characteristics are less likely to have sea pens.

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I am not aware of any freshwater associations with P. gurneyi. My research involved the giant Pacific octopus Jim Cosgrove, Royal BC (Enteroctopus dofleini). Because E. dofleini is a major predator on crabs such as C. productus and C. magister I Museum tended to work in rocky areas that abutted sand or mud flats. Pat Malecha, NOAA Freshwater outflows are not found at the sites. Marine Ecology & Stock Assess In Southeast Alaska fresh water outflows are everywhere…..they do not seem of affect the sea pens. I don’t see Annette Smith, Underwater bigger or smaller forests around fresh water. What I find interesting is why tochuina’s and diomedia’s (both feed on Photographer sea pens) do not comingle. Interesting that you should ask (sounds like a leading question asked by someone who knows!). I am not an Dr. James Murray, ecologist and have not done a careful survey, but it is my impression that sea pen beds are associated with California State Uni freshwater outflows. No freshwater outflows present (nearby = closer than 200 m) where highest aggregations observed. Do see them at John deBoeck, Director of shelf at outlets of tidal but fresh-water-fed inlets/lagoons/bays Dive Industry Asstn of BC None that I can recall Chad King, NOAA

No freshwater outflows observed. Jackie Hildering, Diver

Dr. Verena Tunnicliffe, No freshwater; but P.g. does not venture near the hypoxic zones of Saanich. UVic Prof I was not aware of fresh water or any other unusual abiotic features. They are usually in sandy areas which tend to be relatively featureless except for some species, such as the pen itself, which sticks up above the flat plain. Other Steve Lonhart, NOAA species use that structure for their own purposes (e.g., to hide, to await prey, etc.). Dr. Charles Birkeland, I don’t remember any freshwater outflow at Alki Point or Golden Gardens, but even if there was any fw outflow, the PhD, Prof (Uni Guam lateral distribution and depth distribution would suggest that it was not influential. Marine Lab) Doug Swanston, Diver This species was not observed in Dives on Bowie Seamount Seacology No freshwater outflows on the reefs that I know but seasonal freshwater run off coming form the steep slopes adjacent to the reefs. As the sand allows them to attach to substrate the mud and sand are the main abiotic Peter Mieras, Diver features. do not generally recall fresh water outflows, but do see the sea pens up east side of Vancouver Island off Qualicum, Mike Tonnesen, Diver and out around Hornby Island and inside Denman isl. Melva Nikki van Schyndel, Near a septic tank outpipe and dock. Also in a shallow bay with some freshwater outflows at head. Naturalist in Echo Bay Kingcome and Wakeman Rivers have a strong influence in the environment; the water is often quite brackish and Bill Procter, Broughton brown. Archipelago Dr. Ron Shimek, Prof, UW The pens are common in estuarine situations, but as far as being directly bathed in fresh water, no. Friday Harbour Lab, BMSC etc I don't recall seeing them near any substantial freshwater inputs. They seem to be in areas with good water Dr. Greg Jensen, Uni of circulation - little or no accumulated silt or detritus- but not areas with really strong currents, except juveniles as Washington noted above.

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13 Do you believe that any of the previously-mentioned abiotic (non-living) factors dictate where P. gurneyi choose to aggregate? Please explain. Dr. Chris Harvey-Clarke, Substrate and current seem to be the main factors UBC Yes, I believe that some of the previously-mentioned abiotic factors may dictate where P. gurneyi choose to aggregate. It appears that P. gurneyi require a certain thickness of soft sediment for anchoring themselves against currents, and also to escape from predation, as they will "dig" themselves in as soon as they have been disturbed. Dr. Barb Faggetter, 'Ocean Thick layers of soft sediment are typically found in high sediment environments (e.g., estuaries) in regions of low Ecology' slope and low rugosity. Since P. gurneyi are filter feeders, a moderate to strong current is probably preferred, as this increases the rate at which "food" particles are brought to the organisms by the currents. Dr. David Cowles, Walla Sandy substrate is very important. I know they can live in quieter waters as well. Walla Uni Prof Silty sand sediments are obviously the preferred sediment type for P. gurneyi, but because individuals and low- Michael Kyte, Senior density populations occur in a variety of sediment types (e.g., silty gravel pockets in rocky habitats), this not an Marine Biologist absolute habitat controlling factor. Dr. Dennis Willows, Uni of Yes, sand, mud, current, and freshwater seem to be correlates. Washington Prof Dr. Gary Williams, PhD, N/A Curator Invert Zoology Absolutely. I do not know how the settlement behaviors produce it, but the patchy distribution clearly suggests some factors might be favorable for targeting settlement. I doubt flow alone, because there are many sites with Dr. Russell Wyeth, St FX similar flow and inappropriate substrate. Thus, I would hypothesize substrate cues, conspecific cues or community- asst. Prof member cues might all contribute, possibly with the addition of flow cues. Yes. Because P. gurneyi tends to be a non-motile filter feeded the availability of nutrient rich currents are essential. Jim Cosgrove, Royal BC Colonies are smaller and more dispursed as current levels drop. Museum Pat Malecha, NOAA Substrate is likely a predictive variable for P. gurneyi aggregations. Marine Ecology & Stock Assess Here they live in silty bottoms. I never see them on rocky bottoms or on walls. They tend to like gentle slopes and Annette Smith, Underwater silt! Photographer I’d love to hear your final report on this question! I’ll just say again, I do believe they tend to aggregate around Dr. James Murray, freshwater inflows. California State Uni Yes- at least some current flow seems to be required to thrive. I would say ‘lack of siltation’ may also be a John deBoeck, Director of ‘preferred’ environment, given highest ‘observed’ concentrations have been away from (as in not within the silty Dive Industry Asstn of BC waters) from heavy mineral siltation I do not have enough information to answer this. Chad King, NOAA

Sand, low flow, shallow depth, high shell content. Jackie Hildering, Diver

Dr. Verena Tunnicliffe, It’s likely they are not tolerant of low oxygen UVic Prof

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I have never seen them on rocks. I believe they are limited to sandy habitats, but not sure if there is a limit due to Steve Lonhart, NOAA grain size. I am not aware of other abiotic factors that may influence its distribution. Dr. Charles Birkeland, No. See answer 12. PhD, Prof (Uni Guam Marine Lab) Doug Swanston, Diver Current Seacology See my previous comments Peter Mieras, Diver

No...flatter sandy soft bottoms seem to be where they like to aggregate. Mike Tonnesen, Diver

Melva Nikki van Schyndel, No Naturalist in Echo Bay Bill Procter, Broughton Unsure Archipelago Dr. Ron Shimek, Prof, UW Yes, the sediment is of paramount importance. The major factors are probably sediment particle size distribution Friday Harbour Lab, BMSC and percent organic matter. etc Yes, I suspect they don't handle low salinity very well. Areas with very low currents probably don't deliver enough Dr. Greg Jensen, Uni of food to the colonies, and those with really strong currents probably uproot them. Washington Are you aware of, or have any hypotheses regarding additional abiotic (non-living) environmental characteristics that may control P. gurneyi 14 distribution and fitness? Dr. Chris Harvey-Clarke, No UBC P. gurneyi apparently feed primarily on phytoplankton; their bright orange colour is the result of carotenoids derived from a diet of dinoflagellates [Best, B.A. 1988. Passive suspension feeding in a sea pen: effects of ambient flow on volume flow rate and filtering efficiency. Biol. Bull. 175:332-342.]. Since the majority of large P. gurneyi Dr. Barb Faggetter, 'Ocean aggregations that I have seen have been in the euphotic (sun-lit) zone from 10 m to 30 m, I would suggest that light, Ecology' which is required for phytoplankton growth, may also be an additional abiotic factor that may control P. gurneyi distribution and fitness. Dr. David Cowles, Walla No Walla Uni Prof I am familiar with most if not all of the literature on P. gurneyi, and know of no hypotheses or theory about Michael Kyte, Senior environmental characteristics that control P. gurneyi other than sediment type and adequate water flow with good Marine Biologist plankton resources. Dr. Dennis Willows, Uni of No Washington Prof Dr. Gary Williams, PhD, An appreciable current or water movement is necessary. Curator Invert Zoology

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I would hypothesize that substrate disturbance, suspended sediment, and large salinity fluctuations would all be detrimental to P. gurneyi. I would hypothesize that nutrient input, in so far as it leads to increased plankton might Dr. Russell Wyeth, St FX benefit P. gurneyi (but only if nutrient load is low enough and flow high enough that the substrate does not begin to asst. Prof support fungal colonization or even anoxia). Anything affecting Hippasterias distributions, and possibly Armina distributions would also likely have substantial conseqences on P. gurneyi. Tritonia also, but less so. I am unaware of the literature related to P. gurneyi and its environmental requirements such as salinity, Jim Cosgrove, Royal BC contaminants, etc. but any change in those factors that moves the environment away from optimal will have a Museum negative effect on the fitness and distribution of the colony. Pat Malecha, NOAA No Marine Ecology & Stock Assess Annette Smith, Underwater No Photographer Yes, I hypothesize that they are found in shallower water because they depend on higher currents. At greater Dr. James Murray, depth there is lower current levels, and perhaps less plankton. Maybe this is a reason why sea pens are not found California State Uni deeper and are supplanted by sea whips. John deBoeck, Director of No Dive Industry Asstn of BC I do not have enough information to answer this. Chad King, NOAA

Hypothesis: Sand and high shell content substrate and low flow. Often shallow water. Jackie Hildering, Diver

Dr. Verena Tunnicliffe, No UVic Prof I would think grain size and flow would be important, since the feeding structure is relatively fragile and could not Steve Lonhart, NOAA withstand high turbidity loads and water forces that could scour soft tissues. Dr. Charles Birkeland, I am unaware of any. PhD, Prof (Uni Guam Marine Lab) Doug Swanston, Diver Able to proliferate in waters with periodic significant fine particulates loading. Seacology Possibly pollution in the substrate may diminish density or presence of the orange seapen Peter Mieras, Diver

No Mike Tonnesen, Diver

Melva Nikki van Schyndel, No Naturalist in Echo Bay Bill Procter, Broughton No Archipelago

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Dr. Ron Shimek, Prof, UW Current flows and sediment types are intimately intertwined in helping determine the distribution of the pens. Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of None come to mind Washington PART C. ASCRIBED VALUE 15 Do P. gurneyi (or sea pens in general) hold any specific traditional or cultural role for you? Please explain. They have an intrinsic role as part of the overall marine biodiversity here in the PNW. As well as a giant species Dr. Chris Harvey-Clarke, they form part of the general trend to giantism seen in some PNW taxa which I think helps support their uniqueness UBC and special ecological value. Dr. Barb Faggetter, 'Ocean No Ecology' Dr. Chris Harley, UBC No Associate Prof Dr. David Cowles, Walla Not traditional or cultural Walla Uni Prof Michael Kyte, Senior No Marine Biologist Dr. Dennis Willows, Uni of No Washington Prof The animals I study (primarily Tritonia) depend critically on sea pens as prey. I would have to change my research Dr. Russell Wyeth, St FX focus if sea pen beds disappeared. asst. Prof Jim Cosgrove, Royal BC No Museum Pat Malecha, NOAA No Marine Ecology & Stock Assess No. Other than they are quite beautiful to see underwater. They are also awesome at night when you can watch Annette Smith, Underwater them sparkle if you run your fingers up them. Photographer Dr. James Murray, No cultural value per se, but they are beautiful and a sea pen forest is a fantastic place to visit! California State Uni Yes- largest octo-coral / soft coral species in B.C. and UW photographers love them John deBoeck, Director of They ‘bio-luminesce’ when stroked gently at night Dive Industry Asstn of BC No Chad King, NOAA

No Jackie Hildering, Diver

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Dr. Verena Tunnicliffe, No UVic Prof No Steve Lonhart, NOAA

Dr. Charles Birkeland, No PhD, Prof (Uni Guam Marine Lab) No Dr. David Arsenault, BMSC

Doug Swanston, Diver Beautiful organism Seacology The orange sea pen is an attractive part of the entire marine ecosystem that supports our business. Divers and Peter Mieras, Diver underwater photographers pay money to dive and see them. No, just enjoy and observe, always enjoyed by sport divers and photographers Mike Tonnesen, Diver

Melva Nikki van Schyndel, No Naturalist in Echo Bay Bill Procter, Broughton They are unique and very beautiful and rare. Archipelago Dr. Ron Shimek, Prof, UW Yes, they are one of the neatest animals around. Cool critters. Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of No Washington Is it important for you to know that P. gurneyi (or sea pens in general) simply exist, even if they do not play a role in your culture/lifestyle (ie. 16 existence value)? Please explain. Dr. Chris Harvey-Clarke, Yes, see above UBC Yes. Biological diversity and natural habitats have high intrinsic value for me. Dr. Michael Hart, SFU

Yes. I believe all organisms have important roles to play in the ecosystem, whether we understand those roles or Dr. Barb Faggetter, 'Ocean not. Therefore, it is important to me that all organisms exist and create the complex web of life that is a part of our Ecology' ecosphere. Dr. Chris Harley, UBC Yes. They are very unique animals. And beautiful. Associate Prof Dr. David Cowles, Walla Yes Walla Uni Prof

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Yes it is important to me to know that P. gurneyi and sea pens in general exist. I am a professional zoologist, marine biologist, and natural historian. Sea pens and other invertebrates have been a central part of my life since I Michael Kyte, Senior was a young teen-ager. These organisms are an essential part of my environment that I enjoy directly when I visit Marine Biologist the shore and vicariously through various media. Indeed, I have chosen the geographical area in which I live because of the presence of such organisms as sea pens. Dr. Dennis Willows, Uni of Yes because they are sole food source for nudibranchs that are important in my research. Washington Prof Dr. Russell Wyeth, St FX Absolutely yes. I appreciated and value all of nature’s diversity. asst. Prof All life has value. The loss of P. gurneyi would likely result in negative impacts for nudibranch. One has only to look at all the species that have gone extinct in the half billion years whether due to natural acts or due to human acts. How ironic is it that the state flag of California is the Golden Bear? Humans drove that species extinct. Now Jim Cosgrove, Royal BC we spend millions and millions of dollars to conserve endangered species of no more consequence than P. gurneyi. Museum Why would we even consider an action that would cause a species to go extinct regardless of what the benefits might be? Pat Malecha, NOAA Yes, in the sense that they are part of the benthic ecosystem. Marine Ecology & Stock Assess Yes. Sea pens provide habitat for other critters. They also provide food for other critters. Tochuinas that live in the Annette Smith, Underwater sea pen forests are huge compared to the tochuinas you see elsewhere. Photographer Yes, their basic body form has been in the fossil record for 500 million years. They are one of the first successful Dr. James Murray, animal types. California State Uni Perhaps for purposes of ecosystem stasis. Chad King, NOAA

Yes! I believe that they are important in the marine ecosystem, representing very significant biomass and being a key prey item for multiple species e.g. the sea slug species I have mentioned. Likely you have seen this summation of research? http://reefkeeping.com/issues/2005-08/rs/feature/index.php I Jackie Hildering, Diver very much believe too “ In many respects, Ptilosarcus gurneyi in sea pen beds fill an ecological position similar to the huge herds of bison that used to occupy the American Great Plains, or to the grazing animals of the Serengeti. In each of these cases, whole food webs were built upon the basic keystone resource species.” They are likely significant as ecological engineers and in the provision of ecosystem functions such as flow Dr. Verena Tunnicliffe, modification and shelter UVic Prof Yes. I am a marine ecologist working for the Federal government, and understand these organisms play a role in the benthic, soft-sediment community. It is a shame they are so poorly understood, but that does not make them Steve Lonhart, NOAA unimportant. Dr. Charles Birkeland, Yes. PhD, Prof (Uni Guam Marine Lab) Yes, as I am a strong believer in maintaining biodiversity Dr. David Arsenault, BMSC

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Doug Swanston, Diver It is important to respect the habitat created by P. gurneyi as well as the locations were these organisms exist. Seacology YES see my previous answer Peter Mieras, Diver

one of the more colorful things to see while diving, along with the variety of nudibranchs Mike Tonnesen, Diver

Yes, I often try to show people sea pens on my encounter tours here in Echo Bay as they really capture the Melva Nikki van Schyndel, imagination and their life story is quite fascinating. Naturalist in Echo Bay Bill Procter, Broughton Yes, see above Archipelago Yes. These are beautiful organisms and a favorite for recreational divers in particular. Dane Stabel, Diver

Dr. Ron Shimek, Prof, UW Of course. Friday Harbour Lab, BMSC etc Dr. Greg Jensen, Uni of Yes- they're interesting organisms themselves and support an interesting assemblage of other animals. Washington Are you aware of, or have any hypotheses regarding important ecosystem functions (physical, chemical, and biological processes/attributes 17 that contribute to the ecosystem) attributable to aggregations of P. gurneyi? Dr. Chris Harvey-Clarke, Interest in chemical signaling and bioactive molecules in cnidaria including this species UBC Large individuals of P. gurneyi can filter over 100,000 particles/s. A dense bed of P. gurneyi could have significant impacts on populations of phytoplankton and zooplankton in the region over the bed. Thus, they may have some Dr. Barb Faggetter, 'Ocean local impact over phytoplankton and zooplankton population densities through "bottom-up" control. Additionally, Ecology' dense beds of P. gurneyi provide food for other organisms (nudibranchs and sea stars, for example), as well as refuge/habitat for some commercial organisms (Dungeness crab, rockfish). A general theory that should apply is that of ecosystem engineering. In other words, sea pens provide habitat Dr. Chris Harley, UBC structure, both above and below the sediment surface. Associate Prof Dr. David Cowles, Walla P. gurneyi definitely provide structure to the ecosystem they inhabit, plus are a source of food for several predators. Walla Uni Prof As remarked on by Dr. Ron Shimek (see Question 18) and me (2001), a dense aggregation of P. gurneyi is a biomass-rich environment that as a “keystone” species supports a diverse group of predators. In addition, an aggregation of P. gurneyi provides a relatively “rough” texture environment that provides shelter to another group of species (e.g., juvenile rockfish and flatfish). Another aspect is that a dense aggregation of P. gurneyi will affect water flow velocities at the water-sediment interface modifying the benthic environment. However, none of these Michael Kyte, Senior factors have been published or written in the form of a formal hypothesis or theory regarding ecosystem functions. Marine Biologist Indeed, sea pens have been noted as large Infauna but otherwise largely ignored in the marine biological scientific literature except for their taxonomy. Dr. Ron Shimek probably has described this situation best by his statement “This species is as typical of the Pacific Northwest as apples and sasquatch, and yet it is almost as unknown as the giant Palouse earthworm. And that is a pity!” I fully agree with his sentiment.

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Dr. Dennis Willows, Uni of No Washington Prof No, but this is solely due to lack of attention. I can only think of Metridium anenomes as candidate for a suspension Dr. Russell Wyeth, St FX feeder with larger planktonic predation. Given sea pen densities in some locations I would hypothesize that have asst. Prof relatively strong effects on plankton and thus ramifications through marine food webs in those locations. This is not my area of research but I am willing to speculate that the loss of P. gurneyi from a habitat would also Jim Cosgrove, Royal BC result in the loss of other species. Museum Pat Malecha, NOAA I do not know of their ecosystem function but they likely provide habitat, in the form of vertical structure, for fish, and Marine Ecology & Stock invertebrates. Assess Annette Smith, Underwater I am not a scientist…..just a diver who has been diving in the area for years. Photographer I assume that since they eat plankton, they are important in nutrient cycling. But as they are likely toxic to most Dr. James Murray, species, they directly support only a few species of predators and do not provide habitat to many other species as California State Uni does eel grass. No Chad King, NOAA

Very significant biomass making the energy of plankton available to the species’ predators. Behaviour of Jackie Hildering, Diver “expanding and contracting” several times a day may have impact on the substrate for other organisms. They are likely significant as ecological engineers and in the provision of ecosystem functions such as flow Dr. Verena Tunnicliffe, modification and shelter UVic Prof Almost certainly they provide biogenic habitat. Steve Lonhart, NOAA

Dr. Charles Birkeland, No PhD, Prof (Uni Guam Marine Lab) No Dr. David Arsenault, BMSC

They sustain important food webs. Some of the organisms in the food web provide import functions for humans Doug Swanston, Diver including providing value to a diverse range of human endeavors including Neurobiology Seacology No but I assume if you ask a researcher that specializes in this species may help you Peter Mieras, Diver

No Mike Tonnesen, Diver

Melva Nikki van Schyndel, No Naturalist in Echo Bay Bill Procter, Broughton I think that they filter things out of the water. Archipelago Dr. Ron Shimek, Prof, UW I presume that they drastically influence current flow over the substrate. I have already mentioned their bioturbatory Friday Harbour Lab, BMSC effects. etc I suspect dense beds (when extended) have a significant effect on water flow close to the substrate, which could Dr. Greg Jensen, Uni of influence settling invertebrate larvae. They could also be important predators on invert larvae, including larval Washington geoducks.

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20 Is there any other information regarding P. gurneyi that you would like to add? Please explain. Sea pens have been declining in Puget Sound. Some researchers who used to work on them there (e.g., near Dr. Chris Harley, UBC Golden Gardens State Park in Seattle) can’t find them in such abundance, if at all. Not sure if that is also true in the Associate Prof Strait of Georgia, but it does at least suggest that this species might be one of conservation concern. It is notable that Birkeland (1968:10) stated “In general, then, Ptilosarcus is never sparse.” This statement was verified by Birkeland at a number of locations in Puget Sound. Thus, it is somewhat alarming that since approximately 1980, that I have found the P. gurneyi populations in all of Birkeland’s original study areas to be sparse and only a relatively small fraction of their original density (Kyte 2001). However, the population that I Michael Kyte, Senior recently observed on the west side of Ketron Island in South Puget Sound appeared to be nearly as dense as those Marine Biologist studied by Birkeland. The only other location that may have dense P. gurneyi populations is Dash Point near Tacoma, Washington. This location is near the entrance to South Puget Sound. Wyeth et al. (2006) observed the P. gurneyi predator Tritonia diomedea at this location and their photographic material shows a relatively dense P. gurneyi population. I have a number of tables and maps indicating both possible and known sites in the San Juan Islands and Puget Sound. Dr. Russell Wyeth, St FX I would very much like to see a report, particularly something gives a summary of known sea pen bed locations. It asst. Prof would be invaluable in my research. You may like to contact Mr. Kelly Sendall (Manager of Natural History) at the Royal British Columbia Museum at Jim Cosgrove, Royal BC (250) 387-3544 and ask him for a copy of all the Museum’s data for P. gurneyi. They may have some interesting Museum data for you. Annette Smith, Underwater I would be delighted to dive areas with someone, if they wanted to see our sea pen forests. Photographer They do seem quite resilient to normal predation and the Tacoma WA site was very near a SuperFund site and yet Dr. James Murray, the pens seemed quite healthy. California State Uni I have emailed a map of observations and the corresponding spreadsheets, which entail depth, lat/long, and in Chad King, NOAA some cases, links to videos or photos. **SEE SERVER It is really important to realize how little is known about the species, how important its biomass is, that it seems to be a key prey item to multiple species, that it is often in the shallows where it is particularly susceptible to the impacts of urbanization (chemical, physical disturbance), and – that research indicates in needs low flow areas Jackie Hildering, Diver whereby any development impacting flow is highly likely to have an impact (again particular susceptibility in the shallows). For an approach to studying fish/coral relationships, you may be interested in: Dr. Verena Tunnicliffe, Du Preez, C., and V. Tunnicliffe. 2011. Shortspine thornyhead and rockfish (Scorpaenidae) distribution in response UVic Prof to substratum, biological structures, and trawling. Marine Ecology Progress Series, 425: 217-231. Dr. Charles Birkeland, It is all in Birkeland 1974. You cited it so I assume you have a copy. If you need a copy, let me know. PhD, Prof (Uni Guam Marine Lab)

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do spend about 60 to 70 days in the water in lots of areas of the coast, May to Sept, at Langara, and March in the French Creek area north to Cape Lazo, spawn herring ass. and Oct, cuke harvesting Campbell River north, Nov, Mike Tonnesen, Diver Dec in Victoria Sidney Dr. Ron Shimek, Prof, UW My consulting rate is US$100/hour; maybe you should hire me for a few (dozen J ) hours Friday Harbour Lab, BMSC etc Beds don't seem to be a stable feature. They seem to persist for a couple decades and eventually get wiped out by Dr. Greg Jensen, Uni of predators; in the meantime new beds pop up (and many blink out while still juveniles) but some eventually take Washington hold. I think many of the juvenile beds succumb to non-specialist predators like Hermissenda.