Biodiversity and the Future of the Gulf of Maine Area Lewis Incze and Peter Lawton Genes

Biodiversity is the diversity of life at all levels of organization, from genes to species, communities and ecosystems.

Species

Nearshore Offshore Bank Basin Slope GoMA: Ecosystem Field Project Habitats and Seamount Communities Abyssal Plain From microbes to whales, and from fundamental biodiversity to EBM GoMA Areas of Work:

Species in the Gulf of Maine Area Ecology: past and present Technology Synthesizing Knowledge Linkages to EBM Outreach

Today’s Agenda:

08:45-09:45 Presentation: The Global Census and GoMA: What did we do? What did we learn?

09:45-10:00 Q&A

10:00-10:20 BREAK

10:20-11:00 Presentation: Pathways to EBM

11:00-11:45 Discussion

Programs of the Census of Marine Life

ArCoD Arctic CMarZ Zooplankton CAML Antarctic Creefs Coral Reefs CenSeam Seamounts GoMA Gulf of Maine Area CheSS Chemosynthetic Systems ICOMM Microbes COMARGE Continental margins MAR-ECO Mid-Ocean Ridges CeDAMAR Abyssal Plains NaGISA Intertidal/Shallow Subtidal CenSeam Seamounts TOPP Top Predators

HMAP History of Marine Animal Populations FMAP Future of “ “ “ OBIS Ocean Biogeographic Information System

Collaborators/Affiliated programs Great Barrier Reef Gulf of Mexico BarCode of Life Encyclopedia of Life Oceans film 10 years (2000-2010) 80 countries, 2700 scientists 17 projects, 14 field projects + OBIS, HMAP Xxx cruises, xxxx days at sea, and FMAP ~ $77m leveraged ~ $767 m --need to 5 affiliated projects (field and technology) check 9 national and regional committees >2,500 scientific papers (many covers) books special journal volumes ~1,200 new species identified >1,500 species in waiting Collection in PLoS-ONE, 2010, incl. US and Canadian papers OBIS 28 m records and counting >1 m records in GoMA 43 N

-NAO

Bear Seamount (-1102 m summit)

August SST Who lives in the Gulf of Maine Area?

GoMRMS WHAT TYPE OF SYSTEM IS THIS? System comparisons, insights Source of Information

Atlantic Reference Center, Huntsman Biological Station Canadian RMS Program, DFO Board of Editors

GoMODP => NECODP (geospatial data standards, OBIS format) Ocean Observing Northeast Region Ocean Council

OBIS =>IOC Now >1 m records in GoMA [US and Canadian demersal trawl and other LMR surveys; NOAA Benthic; US EPA; early expeditions (Albatross); museum] Evolving utilities

New Discoveries New species Range extensions Habitat discoveries Genetic composition and dispersal Patterns of Diversity

Coleman Rarefaction Curves, Fish: Fall NEFSC Bottom Trawls, 1963 – 2008 (45 years) Total species in Surveys = 212 90 (demersal)

80 Continental Slope Georges Bank Expected # species/100 trawls

70 Southern Coastal Shelf

Central Gulf of Maine 60 Northern Coastal Shelf 3 Basins 50 Scotian Shelf Eastern Coastal Shelf 40 Scotian Shelf Browns Bank Browns Bank Expected No. No. ofExpected Species Eastern Coastal Shelf 30 Northern Coastal Shelf Southern Coastal Shelf Expected Number of Species Rarefaction) (Coleman of Number Expected Georges Bank Georges Basin 20 Jordan Basin Wilkinson Basin Central Gulf Continental Slope 10 0 20 40 60 80 100 120 140 No. Numberof Bottom of Samples (Bottom Trawls Trawls) Who lives in the Gulf of Maine? Viruses 106 Archaea/Bacteria

105

Meiofauna ? 3000 unknown 2000 known (~4K )

Species Species Types 1000

10-8 10-6 10-4 10-2 10 0 Length (m)

“monitored” “managed”

10-2 10 0 Knowledge Base “Role”

Viruses Function Supporting 6 10 Composition Regulating Archaea/Bacteria Structure

105

Meiofauna Function, Composition ? 3000 unknown 2000 Microalgae known (~4K ) Composition Provisioning

Species Species Types 1000 Structure Cultural Function 10-8 10-6 10-4 10-2 10 0 Length (m)

“monitored” “managed”

10-2 10 0 Condensed Biodiversity Table from GoMA

# Species # Types Viruses ------Bacteria/Archaea 23 est. 105-106 * Protozoa 41 Chromista 831 ** Dinoflagellata 147 bacteria Fungi 1 Plantae --- Invertebrates 3,519 phytolankton Vertebrates Pisces 865 Reptilia 5 Aves 193 Mammalia 34 Insecta 10

TOTAL 5,669 * W. Li et al. PLoS-One Submitted Dec. 2010 ** Fig. 6 from Li et al. GoMA Field Studies: Breadth and Depth

• Sponsored and collaborative projects

• Ranging across many spatial scales and considerations of system complexity

• Sampling Gulf of Maine habitats from intertidal to 3000 m depths

• Application of a wide range of technologies – “from rubber boots to ROV’s”

• How to organize, represent , and apply the new knowledge stemming from these studies? Genes Invasive species trouble spots: tracking the spread of European green crabs in the Gulf of Maine (J.E. Beyers and J. Pringle, U. New Hampshire; J. Roman, U. Vermont). Genetic analysis from 30 sites on speed of movements of a recently invaded haplotype (on the CO1 gene) from northern Nova Scotia.

Cashes Ledge: system change 20 years later (J. Witman, Brown U.). Tethered predation experiments and time-lapse photography.

Species Platts Bank: part of a network of feeding hotspots Nearshore for upper trophic level predators in the Gulf of Bank Offshore Maine (L. Incze, U. of Southern Maine; S. Kraus, N. Basin Slope E. Aquarium; and D. Fields, Bigelow Lab. (with

Habitats and Seamount collaborations at WHOI and Oregon State U.). Communities Abyssal Plain Nested spatial domains of the Gulf of Maine ecosystem

Closely linked to marine habitat mapping approaches, such as the Gulf of Maine Mapping Initiative

Broader conceptual model needed for spatial analysis, considering connectivity :

• biological processes (larval dispersal, adult migration, benthic-pelagic coupling)

• ocean processes (transport

and mixing) Scales of species interactions within communities and habitat types (first two domains) Species and Communities: Habitat-scale investigations Natural Geography of Inshore Areas (NaGISA **): Cobscook Bay (ME) and Simpsons Island (NB).

Cobscook Bay, Maine

Number Not Seen 250 Total Number of Species 200 22 22 150 24 100 18 Some species were observed for the first 50 and only time 165 years ago. (Additional 4 4 1 3 0 2 analyses comparing NaGISA sampling with Cnidaria Ctenophora Plathyhelminthes Nemertea Annelida Arthropoda Mollusca Ectoprocta Echinodermata Number of Species of Number historical records for Cobscook Bay: T. Trott and C. Kulfan, Suffolk U.).

** NaGISA Regional Coordination: Phylum G. Pohle, Huntsman Marine Science Centre Two perspectives on intertidal species diversity in Cobscook Bay

Gulf of Maine Species Richness Versus Latitude West Atlantic/East Atlantic

1000 1000

Cobscook Bay Cobscook Bay Dale Roads, Wales 800 800

600 600 Casco Bay Mt Desert Region Mt. Desert Region Casco Bay

400 400 SpeciesRichness Numberfo Species Casco Bay Penobscot Bay Penobscot Bay 200 200 Watwick Bay Sheepscot Sheepscot Sheepscot Lydstep Haven Sheepscot 0 0 42 44 46 48 50 52 54 43.4 43.6 43.8 44.0 44.2 44.4 44.6 44.8 Latitude Latitude

T. Trott, Suffolk U. Bank-scale views: Ocean Acoustic Waveguide Remote Sensing

Synoptic view of schooling herring (Clupea harengus) on the northern edge of Georges Bank, sampled during a 75 s scan (full sweep of red circle) by OAWRS (Ocean Acoustic Waveguide Remote Sensing), 3.4 h after sunset. School densities are −45 (blue) to −33 dB (red) (data from N.C. Makris, Massachusetts Institute of Technology). This is one of a sequence of images showing the formation and movement of herring toward the bank. Bank-scale views: Physical-biological coupling across trophic levels at Platts Bank

L. Incze, U. of Southern Maine; S. Kraus, N. E. Aquarium; and D. Fields, Bigelow Lab. (with collaborations at WHOI and Oregon State U.).

pycnocline euphausiid patches Internal Waves Landscape views: defining selected ocean spaces for exploration and data integration

Stellwagen Bank

Discovery Corridor Stellwagen Bank National Marine Sanctuary : pattern analysis

Increasingly automated techniques for statistical analysis of macrofaunal distributions (S. Gallager, WHOI, and The HabCam Group®).

Stellwagen Bank Ecosystems Services Model: processes analysis (L. Incze, U. Southern Maine; P. Auster, U. Connecticut; L. Kaufman, Boston U., B. Haskell, Stellwagen Bank NMS; and others).

HabCam Images ~1m2/frame Gulf of Maine Discovery Corridor Initiative Four offshore research missions since 2005

Embraced by Canadian Healthy Oceans Network (also applied in Arctic & Pacific)

P. Lawton, E. Kenchington, B. Li (DFO); A. Metaxas (Dalhousie U.); P. Snelgrove, S. Bentley, B. Montevecchi (Memorial U.); C. Lovejoy (U. Laval) Discovery Corridor: Habitat structure investigations

Surprisingly rich suspension-feeding community in Jordan Basin dominated by brittle stars, sponges and sea anemones on a deep (188 m) bedrock ridge, dubbed “The Rock Garden”

Winter skate and deep sea corals in Acoustic coral signatures from ROV mounted multibeam – Canadian Northeast Channel (668 m) Healthy Oceans Network, 2010 Discovery Corridor: New coral species records (2006 mission)

Primnoa resedaeformis Paragorgia arborea New depth and colony size records within the Northeast Channel Coral Conservation Area.

Stauropathes arctica Bathypathes patula New Canadian Maritimes record. New Canadian record established.

(Underwater photo credits: Canadian Scientific Submersible Facility/ropos.com). Gulf of Maine View: Diversity and distribution on slope and seamounts

Spatial distribution of the number of known species across the deep water census area of the Gulf of Maine. Blue dots represent a species record.

Bear Seamount (summit depth 1102 m)

Multibeam bathymetry of GoMA Expert Group: Slope, (clockwise) Bear, Physalia and Seamounts and Other Deep-Water Mytilus seamounts with 10x Environments vertical exaggeration. Lead: N. Kelly; Contributors: E. Shea, A. Metaxas, P. Auster, R. Haedrich, T. Sutton, M. Vecchione M. Vecchione NMFS National Systematics Laboratory photophores on Histioteuthis squid Smithsonian Institution

viperfish, Chauliodus sloani

Bathysaurus agassizi

Bolitaena pygmaea Megalocranchia fisheri Dibranchus tremendus Gulf of Maine View: From surveys to prediction of biodiversity patterns and driving factors CoML Cross-Project Synthesis (with Australian Great Barrier Reef and deep Gulf of Mexico): Influence of 27 physical data layers on biodiversity patterns of benthic invertebrates and demersal fish.

Bathymetry & Derivatives In the Gulf of Maine, Substrate habitat explains about 35% of the variation in Bottom Stress abundance and distribution of the Stratification abundant fish and

Sea Surface invertebrates Temperature

Chlorophyll

Benthic Temperature C. Pitcher *, N. Ellis (CSIRO); P. Lawton *, S. Smith, M. Bottom Salinity Greenlaw, J Sameoto (DFO);

Benthic L. Incze, N. Wolff (U. Southern Irradiance Maine); C. Wei (Texas A&M U.); Bottom Nutrients P. Snelgrove (Memorial U.) * Co-leads Ecosystem

Environmental Genetic Scientific articles processes variability/change Demographic Interpretive reports Over-exploitation processes Interspecific interactions, Ecosystem processes Decision-support tools Habitat degradation Ecological processes & disturbances, Specific indicators Societal concerns Resource use Function Recommendations Management needs

Program Drivers Planning Research Analyses Outputs

Scientific Program Evolution Strong/Buzeta Homarus americanus Cerianthus borealis

Solaster endeca Ciona intestinalis Photo credit: Mike Strong and Maria-Inez Buzeta ACKNOWLEDGMENTS

Lewis Incze Principal Investigator Peter Lawton Co-Principal Investigator Sara Ellis Program Manager Nick Wolff Senior Research Associate Suzy Ryan Education & Outreach Coordinator

SUPPORT STAFF EXPERT GROUP LEADERS Adrienne Adamek Catherine Johnson William Li Chelsie Archibald Noreen Kelly Jeffery Runge Jen Ecker Scott Kraus Michael Sieracki Michelle Greenlaw Peter Larsen Kent Smedbol Ashley Holmes Noreen Kelly Sarah Kirn SENIOR ADVISORS

Michael Fogarty Ron O'Dor Michael Sinclair Robert Stephenson

ADVISORS Jim Luyten Robert Andersen Craig MacDonald Paul Anderson Cameron McLellan Peter Auster David Mountain James Bisagni John Nelson Philip Bogden Thomas Noji John Boreman Judith Pederson Kenneth Brink Gerhard Pohle Ann Bucklin Robert Rangeley Jeremy Collie Evan Richert Mark Costello Andy Rosenberg Lee Dantzler Jeffrey Runge Michelle Dionne Michael Sieracki Kenneth Foote Michael Sissenwine Kenneth Frank Andrew Solow Eugene Gallagher Robert Steneck Joseph Kelley David Townsend Scott Kraus Thomas Trott George Lapointe Lou Van Guelpen Jason Link Michael Vecchione Les Watling Peter Wiebe Charles Yentsch CONTRIBUTORS

Karen Alexander William Leavenworth Peter Auster Kevin MacIsaac Ivar Babb Bruce MacLeod Matthew Bampton Nicholas Makris Sam Bentley Anna Metaxas Mary Black Jim Manning April Blakeslee William Michaels Jeffrey Bolster John Moore Ingrid Brack David Mountain Robert Branton Lauren Mullineaux David Briggs David Phinney Jay Burnett Gerhard Pohle Maria-Ines Buzeta James Pringle Jeb Byers Purnima Raital Stefan Claesson Anna Redden Bruce Collette Joe Roman Andy Cooper Shale Rosen David Fields Andrew Rosenberg Michael Fogarty Jeffrey Runge Scott France Stephen Smith Kenneth Frank Paul Snelgrove John Galbraith Andrew Solow Ruth Gibbons Peter Stevick Glenys Gibson Mike Strong Jonathan Grabowski Thomas Trott Nate Hamilton Page Valentine Karsten Hartel Lou Van Guelpen Katherin Hayhoe Michael Vecchione Martin Jakobsson Les Watling Michael Jech Susan Wigley Kevin Joy Jon Witman Ellen Kenchington Huijie Xue EXPERT GROUP MEMBERS

Robert Anderson Anna Metaxas STUDENTS John Anderson John Neilson Peter Auster Todd O’Brien Adrienne Adamek Ann Bucklin Michael Owen Adam Baukus Maria-Ines Buzeta Andy Pershing Jen Ecker Andrew Cooper Gerhard Pohle Richard Franks Ted Durbin Jooke Robbins Michelle Greenlaw Eugene Gallagher Juliette Rooney-Varga Joe Hodgson Dian Gifford Nancy Shackell Heather McRae Andrew Gilbert Tim Shank Angie Fourgues Walt Golet Liz Shea Ashley Holmes Jonathan Grabowski Rabindra Singh Fred Grassle Peter Stevick POST-DOCS Dick Haedrich Jason Stockwell Jon Hare Tracey Sutton Adrian Jordaan Bob Kenney Lou Van Guelpen Noreen Kelly Kerry Lagueux Mike Vecchione Jason Link Les Watling EDUCATION & OUTREACH Molly Lutcavage Mason Weinrich Jennifer Martin Dave Wildish Verna DeLauer Stormy Mayo Willie Wilson Pam DiBona Gary Melvin Jon Witman Justine Glynn Melissa Wong Peter Taylor Mark Wiley Pathways to EBM

A framework for representing biodiversity knowledge:

• capacity to visualize ramifications of management actions in a complex physical-biological system; • an essential two-directional conduit for communicating ideas, priorities, and findings between the regional science and management communities.

Achieved through considerations of:

• compositional, structural, and functional biodiversity elements for specific ecosystem “compartments”; • biological size spectrum of known and unknown diversity; • nested spatial domains.

But…How do we integrate human society into the picture?

• emerging Ecosystem Services (ES) considerations. Compositional marine biodiversity: identity and variety of biodiversity elements within the system, from genes to the complement of species occurring within particular ecoregions and biogeographic provinces. Structural marine biodiversity: physical organization or pattern of biodiversity Ecosystem occurring within the system including both biotic and abiotic variables that modulate patterns.

Biodiversity conservation policies, management

Genetic strategies, and monitoring processes approaches can be aligned

Demographic with particular elements processes (Lawton et al. submitted) Interspecific interactions, Ecosystem processes

Ecological processes & disturbances, Resource use Functional marine biodiversity elements: Function processes that operate at various spatial, ecological, and evolutionary scales to mold biodiversity composition and structure. Assessing our Knowledge base ?

known (~4K ) Function

GoMA Expert Groups:

Microbial communities Coastal Margins Co-leads: W. Li and M. Sieracki Co-leads: P. Lawton and P. Larsen. Zooplankton and pelagic nekton Benthos & Demersal Nekton Co-leads: C. Johnson and J. Runge Lead: P. Lawton Upper trophic level predators Slope, Seamounts and Other Deep Co-leads: S. Kraus and K. Smedbol Water Environments Lead: N. Kelly Interannual variation in copepod species richness, evenness, and diversity in the Gulf of Maine based on standardized Continuous Plankton Recorder samples. Bars indicate standard error. Mean sample size = 94, s.d. = 38.

Fig. 7 from: C. Johnson et al. Biodiversity and ecosystem function in the Gulf of Maine: pattern and role of zooplankton and pelagic nekton (In revision to PLoS-One: December 2010) Comparison of four regional B-HMAP research programs on marine biodiversity: Relative focus on each biodiversity element within each GoMA program depicted by the GoMx relative size of the three GBRSB ellipses (Based on Ellis et al.: In revision to PLoS-One: December 2010)

Composition Structure Composition Structure Composition Structure Composition Structure

Function Function Function Function GoMA B-HMAP GBRSB GoMx Gulf of Maine Area Census of Marine Life (GoMA), Baltic Sea History of Marine Animal Populations (B-HMAP), Great Barrier Reef Seabed Biodiversity Project (GBRSB), and Gulf of Mexico Biodiversity Assessment (GoMx) Nested spatial domains of the Gulf of Maine ecosystem

Ecological processes and patterns are reciprocally linked

Ecological • Interactions can be Processes scale-dependent, and/or cross-scale

• “Curiosity-driven” science may address all scales and complexities Ecological Patterns • “Issue-driven” science and management application typically considers a subset of spatial domains and factors

Ecosystem Services Geochemical & Climate System Biodiversity Elements Physical disturbance Human System

Specialization CompositionC Ecosystem Services Options/flexibility Species attributes Functional diversity Genes Provisioning $$$ (redundancy) Species and abundance food Communities genes pharmaceuticals Emergent SStructure properties Genes Resiliency Populations Adaptation Habitats Environment (optical, chemical) Communities Variability Meta populations Distributions Cultural $ Connectivity aesthetic Food webs Supporting O production spiritual Dominance/rarity FunctionF 2 recreational

O2 production understanding Processes C fixation [Biotic and abiotic Regulating interactions] Climate feedbacks Bioremediation $ Competition Predation Facilitation Decomposition/Recycling What comes next?

Engagement with the regional ocean management community to help define and support:

• Well-planned programs of continued ocean exploration designed to improve basic knowledge of the GoMA oriented toward biodiversity conservation and integrated ocean area management/EBM:

• Field work on key remaining gaps on compositional, structural and functional biodiversity elements;

• Investments in marine taxonomy capacity and further development of regional species registers ;

• More attention on genetic diversity/cryptic species to understand “hidden” diversity within the system;

• Comprehensive bioinformatics approaches to reinforce biological research & monitoring programs;

• Data submission to Northeast Coastal and Ocean Data Partnership, National Data Management portals, and International Ocean Biogeographic Information System (OBIS) • Further evolution of conceptual constructs as a way forward for scientists and managers to debate and agree on focus points:

• Representation of spatial domains, connectivity, scale-dependent , and cross-scale interactions

• Implementation of Coastal and Marine Spatial Planning:

• Representative Areas and MPA network analysis for inclusion of marine biodiversity conservation criteria; • Ecosystem Services (ES) modeling/field studies (e.g. Northern Mass Bay/Stellwagen Bank, Southwestern New Brunswick Marine Resource Planning Initiative, offshore pilot?)

• Suitability of existing scientific data for ES models and approaches to connect (ES) results to economics and existing laws.

• Recruitment of young “knowledge brokers” - they make an excellent investment in the future and in your agencies Discussion Session Topics:

1. Specific questions on the GoMA presentations (Part 1 or Part 2)?

2. How well does biodiversity conservation fit into your management mandates?

3. Is the relationship between biodiversity (elements) and ES clear?

4. Does the emerging ES framework provide a useful path to EBM?

5. Would ecological studies of ES be significantly convergent with the spatial scales at which CMSP will have to be implemented?