Biological Valuation Workshop Assessing ‘Value’ in marine environments John Roff Acadia University Gent, Belgium, December 2004

What constitutes ‘V alue ’ in marine environments? They are ‘unique’ or ‘valuable’ in some definable way They are ‘unique’ or ‘valuable’ in some definable way They are threatened currently or will be in the immediate future They are threatened currently or will be in the immediate future Public, political or NGO concern is expressed for defined reasons Public, political or NGO concern is expressed for defined reasons There is an opportunity to acquire or designate the area There is an opportunity to acquire or designate the area Priority areas for conservation (PACs) generally have the following attributes/ characteristics or criteria:

I will concentrate largely on the first issue of what constitutes uniqueness and value. Note that what constitutes uniqueness is only one component of what constitutes value. Note that what constitutes uniqueness is only one component of what constitutes value. Unique means that we have only ONE EXAMPLE of some entity. In terms of biodiversity it would therefore be inherently valuable. Unique means that we have only ONE EXAMPLE of some entity. In terms of biodiversity it would therefore be inherently valuable. NOTE that biodiversity is NOT just species diversity. NOTE that biodiversity is NOT just species diversity.

“Value” is a subjective term that can encompass: Uniqueness Uniqueness Other combinations of biodiversity attributes Other combinations of biodiversity attributes Market price Market price Resource supply Resource supply Etc. Etc.

Two criteria for value might be: Value for conservation purposes, i.e. an area has high biodiversity value (e.g. species richness). Value for conservation purposes, i.e. an area has high biodiversity value (e.g. species richness). Value for resource purposes, i.e. an area supplies ecosystem services (e.g. high production of some commercially important shellfish species). Value for resource purposes, i.e. an area supplies ecosystem services (e.g. high production of some commercially important shellfish species). Areas exhibiting these two criteria are frequently separate, e.g. species diversity and productivity are often inversely related (or at least the relationship is parabolic).

Relationship between: species richness and production Number of species Productivity North Sea ? Global ?

Obviously we need to assemble data and map such variables. Here I concentrate on value for conservation purposes. Central question is: Central question is: –What makes some: species/ community/ habitat/ location/area/ ecosystem/ etc. More valuable than another? George Orwell: “All animals are created equal, but some are more equal than others”.

When proposals are made to ‘protect’ certain areas of the marine environment, proponents and activists generally use evocative words to describe their chosen location. They may say that an area requires protection because: It is unique in some way It is unique in some way It is ‘special’ in some way It is ‘special’ in some way It is ‘rare’ in some way (e.g. endangered species/ habitat/ community) It is ‘rare’ in some way (e.g. endangered species/ habitat/ community) It supports economically important resources (fishing grounds/ recruitment area/ nursery/ spawning grounds etc.) It supports economically important resources (fishing grounds/ recruitment area/ nursery/ spawning grounds etc.) It has high biodiversity value (species diversity, or other attribute) It has high biodiversity value (species diversity, or other attribute) It exhibits high ecological or environmental complexity It exhibits high ecological or environmental complexity It exhibits coastal zone aesthetics It exhibits coastal zone aesthetics Rarely are these criteria adequately defined, or quantified.

How do we go about such quantification? We need an inventory to consider at least three sets of elements: The ecological hierarchy and biodiversity framework (e.g. Zacharias and Roff) The ecological hierarchy and biodiversity framework (e.g. Zacharias and Roff) Mapping of ‘Valuable” or ‘Unique’ areas - Distinctive areas Mapping of ‘Valuable” or ‘Unique’ areas - Distinctive areas Mapping of ‘ordinary’ or Representative areas Mapping of ‘ordinary’ or Representative areas This is followed by mapping and analysis of combinations of attributes to yield a framework for decision making.

COMPOSITIONSTRUCTURESPROCESSESGenes Genetic structure Genetic processes Species, Populations Population structure Demographic processes, Life histories Communities Community composition Organism-Habitat relationships Ecosystems Ecosystem structure Physical and Chemical processes After Zacharias and Roff 2000, Cons. Bio. Remind ourselves of the COMPONENTS OF : All the structures and processes that support the diversity of life in the oceans

StructureMutationStructureMigrationStructureSuccessionWatermassCurrents GenotypesDifferentiat.AbundanceDispersion S.Diversity S.DiversityPredationTempTides FitnessDriftDistributRetention S. Richness Competit.SalinityDisturban. Haplotype D Flow Focal Spp Mig/ Drift S. Evenness ParasitismPropertiesGyres Stocks Nat. Select KeystoneGrowthAbundanceMutualismBoundariesRetention Inbreeding Ind. Cond. Reprod.Represent.DiseaseDepth/Pres P-B couple Mating Ind. Comp. RecruitDistinctiveProductionLightEntrain. Dir. Select UmbrellaBiomesDecomp.Stratificat. B-G cycles Stab. select Charismat.Biocoenos.TopographSeasonal. Dis. select Vulnerable S-A relns. SubstrateProduct. Micro. Evol. EconomicTransitionsRepresent. H-A equil. ErosionPhenotypesFun.groupsDistinctive H-L equil. SpeciationFragmentsHeterog.AnomaliesTurbulence Macro. Evol Meta-popsEndemismExposureMixing Alt. S.Stats PatchinessUpwelling SymbiosesNutrientsDivergence Biomass Dis. Gases Ecol.Integ. AnoxiaErosion Desiccation GENETIC Structure Process SPECIES/ POPULATION Structure Process COMMUNITY Structure Process ECOSYSTEM Structure Process The components of marine biodiversity: Expanded from Zacharias and Roff 2000

An entity may be unique or have value - at any level of the ecological hierarchy: - Presence or seasonal use by: Threatened/ rare/ endangered species (flagships?) Threatened/ rare/ endangered species (flagships?) Keystone species (what is it/ does it work?) Keystone species (what is it/ does it work?) Other focal species, feeding breeding areas Other focal species, feeding breeding areas Species of economic value, especially spawning, recruitment, breeding areas Species of economic value, especially spawning, recruitment, breeding areas SPECIES LEVEL:

COMMUNITY/ HABITAT LEVEL: Endangered, rare, threatened habitat Endangered, rare, threatened habitat –i.e. a particular kind of habitat that has been reduced in quantity or degraded to a far greater extent than other marine or coastal habitat types, i.e. the proportion of this habitat in its ‘pristine’ condition is far lower than that of other habitat types. Critical habitat Critical habitat –Equivalent to biologically ‘Distinctive habitats’. Regions of high species diversity (NOT Biodiversity). Regions of high species diversity (NOT Biodiversity). –Often asserted but rarely documented adequately. Do we know WHY some areas have higher species diversity than others? Regions of high topographic complexity or habitat complexity Regions of high topographic complexity or habitat complexity –leading to high species diversity or to locations of rare/ endangered species?

ECOSYSTEM/ HABITAT LEVEL: Regions of unique ecosystem level processes Regions of unique ecosystem level processes –e.g. upwellings, gyral systems, estuarine retention systems. Regions associated with focal species. Regions associated with focal species. These are: Distinctive Habitats These are: Distinctive Habitats

For analysis of habitats, we must consider Structures and Processes across the entire ecological hierarchy For analysis of habitats, we must consider Structures and Processes across the entire ecological hierarchy AND

Processes Processes –Upwelling, Gyres, Currents Environmental Anomaly Environmental Anomaly –Temperature, Topography, Sea Height, Chlor a Focal Species Focal Species –Flagships, Umbrellas, Parasols, Indicators STRUCTURES AND PROCESSES Roff and Evans 2002 Aquat. Cons.

Characteristics, Processes, Focal Species Upwelling areas VentsCoral Reefs Gyres Circulation Seamount Shelf Edge Canyons Sponge Beds Nutrient addition Sulphur bacteria Symbiotic and other algae Physical accumul- ation Sediment flux enhanced ? low diversity high diversity low diversity high diversity Flagships Parasols Indicator species Indicator species Flagship s Parasols ? ? Resources Elevated High Primary Production Areas Retention Areas Resources advected / focused Resources depleted Caves Sediment by-pass ? high diversity Indicators Roff and Evans 2002 Aquat. Cons.

Anomalies and Focal Species - Examples AnomalyLocation Physical Process Focal Species Biological Process Low temp/ high chlor a SW Nova Scotia Upwelling Many larval species Recruitment Cells Topography Saguenay Fjord Estuarine circulation Whales Euphausiids Feeding Topography Islands everywhere Isolation / Geographic SealsBirds Reproduction Feeding Topography / Currents Fundy / P’quoddy Bay Gyre / Tidal Circulation Whales (Copepods) Feeding High temp/ high chlor a Minas Basin Mudflats Tidal Resuspension Migrant Birds Feeding Roff and Evans 2002 Aquat. Cons.

Relations between Anomalies and Focal Species

Bay of Fundy – Grand Manan. Lewin and Roff unpub.

Other Distinctive habitats May not be detected by physical anomalies, satellite data or conventional ocean mapping May not be detected by physical anomalies, satellite data or conventional ocean mapping –Fish spawning areas –Juvenile fish recruitment zones –Turtle breeding beaches –Submarine seamounts –Deep sea vents Located by TEK or SEK Located by TEK or SEK

What about ‘ordinary areas’? The Representative habitats. If we do not protect them, we ignore >90% of marine species diversity! Representative habitats (benthic and pelagic) can be mapped - at some scale Representative habitats (benthic and pelagic) can be mapped - at some scale –we can map areas of high habitat heterogeneity –we can map seasonal and inter-annual variations in geophysical factors – e.g. in water masses –we can determine correspondence between selected assemblages of organisms and geophysical factors (e.g. fish, benthic invertebrates and water masses). –THEN, if there are strong correlations between e.g. water masses and fish assemblages, we can know how these assemblages will change in response – perhaps even in response to global warming. –THEN, we can also potentially locate the limits of distributions of communities, their spatial variations, and their centroids of distribution and abundance.

Map the marine environment to define Representative areas (seascapes) Water Masses Water Masses Temperature Temperature Salinity Salinity Water depth Water depth Water ‘colour’ Water ‘colour’ Currents Currents Substrate types Substrate types Habitat types Habitat types Fish distributions Fish distributions Marine mammals Marine mammals ETC. ETC. ETC. ETC. Water depth ACTUAL SET OF FACTORS USED DEPENDS ON NATURAL VARIATION WITHIN A REGION Roff et al. Aquat. Cons. 2003

ASSEMBLING BENTHIC SEASCAPES For the Gulf of Maine and Scotian Shelf Bottom Water Masses + Depth Classes + Generalized Substrate = BENTHIC SEASCAPES

For the Gulf of Maine and Scotian Shelf Notes: ~100 Benthic Seascape Types Each Type defined by its components Unique Code Identifies each # ,000 (Water Mass 6 – Maine SW) 200 (depth m) 40 (D - gravel/till) __________ 6240

ASSEMBLING PELAGIC SEASCAPES For the Gulf of Maine and Scotian Shelf Surface Water Masses + Depth Classes + Stratification Classes = PELAGIC SEASCAPES

For the Gulf of Maine and Scotian Shelf Notes: ~70 Pelagic Seascapes Each Type defined by its components Unique Code Identifies each # ,000 (water mass 25) -200 (depth ) -20 (frontal) __________

Scotian Shelf Seascapes of Representative Habitats Roff et al Aquat. Cons.

Habitat heterogeneity How MANY different kinds of Representative Habitats? Roff et al Aquat. Cons. What can we do with these seascapes?

T S Temperature – Salinity water masses

Scotian Shelf / Gulf of Maine – bottom Water Masses (variability)

CORRESPONDENCE BETWEEN FISH COMMUNITIES AND OCEANOGRAPHY COURTESY KEES ZWANENBURGCOURTESY WWF / CLF WATER MASSES T-S FISH COMMUNITIES

Relation between benthic invertebrate assemblages and water masses (Roff and Lewin unpublished)

Centroids of benthic community distributions (Lewin and Roff unpublished)

Representative habitats are NOT unique. BUT: Do some have more ‘value’ than others? Are all replicates of representative habitats created equal? Apparently not, but we do not yet know all the ‘rules’. We know that: Different habitats have different communities that vary in their species diversities. Different habitats have different communities that vary in their species diversities. –It may be interesting scientifically to ask why this should be so. However, for conservation purposes we should simply accept it, and try to capture all types of representative communities. It is WITHIN habitat and community types that we need to ask what species diversity is related to. It is WITHIN habitat and community types that we need to ask what species diversity is related to.

Zacharias and Roff. Innes and Roff unpublished Temperature, Salinity, physical & biological disturbance effects. Temperature, Salinity, physical & biological disturbance effects. Relationships between biomass and species diversity. Relationships between biomass and species diversity. Confounding effects of: current speed (in bringing resources, causing disturbance). Confounding effects of: current speed (in bringing resources, causing disturbance). Hypotheses being tested. Hypotheses being tested. Biomass increases with POC flux. Biomass increases with POC flux. Species richness Species richness –increases with physical disturbance (fetch), –decreases with temperature and salinity fluctuations, and –increases with habitat complexity.

BC Intertidal Data set 370 station by 500 species intertidal data set from British Columbia Collected during summer daylight low tides Collected during summer daylight low tides Standardized sampling methodology Standardized sampling methodology Qualitative abundance Qualitative abundance Macrobiota (no rock turning) Macrobiota (no rock turning) No fine sediment beaches No fine sediment beaches Slope, aspect, substrate, salinity, temperature, fetch, precipitation Slope, aspect, substrate, salinity, temperature, fetch, precipitation Zacharias, Roff & Howes (2001) J. Biogeog. SPECIES DIVERSITY – WITHIN REPRESENTATIVE HABITAT TYPES

Diversity at the Species / Habitat Level Salinity versus Species Numbers

Fetch versus species numbers Fetch Number of species

R 2 = 0.71 Diversity at the Species / Habitat Level Combined effects of: Salinity, Temperature and Fetch PHYSIOLOGICAL disturbance REDUCES species diversity PHYSICAL disturbance INCREASES species diversity

Predators versus other species When Predator effects are added: R 2 = 0.86 BIOLOGICAL disturbance increases diversity

Current research - Representative habitats Innes – sub-tidal communities Innes – sub-tidal communities –Relation of biomass to resource flux/ production –Topography, complexity, disturbance – relation to species diversity –Relation of species diversity to resource flux Lewin - Scotian Shelf zoobenthos Lewin - Scotian Shelf zoobenthos –Define habitat-community associations –Define species-area curves

Essentially we are asking: what are the surrogates for species richness of a region? ANSWER: This is not well known! HOTSPOT paradigm. Presence of higher taxonomic level members –e.g. birds? Does not seem to work. In general the presence of such ‘charismatic mega-fauna indicators’ simply means that a region is rich in resources at some trophic level. This usually means that such areas are in fact LOW in species diversity – at least at some trophic level. HOTSPOT paradigm. Presence of higher taxonomic level members –e.g. birds? Does not seem to work. In general the presence of such ‘charismatic mega-fauna indicators’ simply means that a region is rich in resources at some trophic level. This usually means that such areas are in fact LOW in species diversity – at least at some trophic level. BENTHIC COMPLEXITY. Not yet well tested – Innes work. BENTHIC COMPLEXITY. Not yet well tested – Innes work. RICHNESS, VULNERABILITY, RARITY? RICHNESS, VULNERABILITY, RARITY?

Other ‘notions’ ECOLOGICAL INTEGRITY: ECOLOGICAL INTEGRITY: –I am not a ‘fan’ of this notion. Essentially there is no such thing as ecological integrity of a single marine habitat. It depends critically on patterns of connectivity in the region not on isolates. NATURALNESS: Two major problems: NATURALNESS: Two major problems: –How do we know what the natural state of an area should be? –Shifting baseline problem –Potential for indicators of community composition and condition – not well developed for marine communities. PRODUCTIVITY: PRODUCTIVITY: –At what trophic level? –Is it inversely related to species diversity? –Is it not an asset for commercial fisheries? I.e. these may be the very areas that we do NOT want to conserve?

Most important areas to protect (Priority Areas for Conservation) are: Areas that are truly unique (one of a kind) Distinctive Habitats Areas that are truly unique (one of a kind) Distinctive Habitats Examples of each type of Representative Habitat – to capture the greatest range of species diversity. Examples of each type of Representative Habitat – to capture the greatest range of species diversity. Areas that have ‘added value’ – where several components of biodiversity coincide or spatially overlap i.e. COMBINATIONS of Distinctive and Representative Habitats. Areas that have ‘added value’ – where several components of biodiversity coincide or spatially overlap i.e. COMBINATIONS of Distinctive and Representative Habitats. Areas that would protect or conserve economically valuable renewable resources. Areas that would protect or conserve economically valuable renewable resources.

From inventory and mapping exercises, we can now plan so as to capture as many components of biodiversity as possible within a region. We can see how to make decisions using e.g. MARXAN analysis. We can select: Areas that have the greatest ‘ecological value’ (greatest ‘bang-for’ the-buck’) Areas that have the greatest ‘ecological value’ (greatest ‘bang-for’ the-buck’) Areas that combine the greatest number of features from our inventory in the ecological hierarchy of biodiversity. These are in fact now: Areas of high biodiversity value (is this not what we are after?) Areas that combine the greatest number of features from our inventory in the ecological hierarchy of biodiversity. These are in fact now: Areas of high biodiversity value (is this not what we are after?)

MARXAN ANALYSIS All Species / Years Data Irreplaceability Results BLM = 5 Target = 25% Sydney Bight Haddock Box Halifax Area Mouth of BOF SW NS Upwelling Evans and Roff unpublished COMBINING REPRESENTATIVE AND DISTINCTIVE AREAS

StructureMutationStructureMigrationStructureSuccessionWatermassCurrents GenotypesDifferentiat.AbundanceDispersion S.Diversity S.DiversityPredationTempTides FitnessDriftDistributRetention S. Richness Competit.SalinityDisturban. Haplotype D Flow Focal Spp Mig/ Drift S. Evenness ParasitismPropertiesGyres Stocks Nat. Select KeystoneGrowthAbundanceMutualismBoundariesRetention Inbreeding Ind. Cond. Reprod.Represent.DiseaseDepth/Pres P-B couple Mating Ind. Comp. RecruitDistinctiveProductionLightEntrain. Dir. Select UmbrellaBiomesDecomp.Stratificat. B-G cycles Stab. select Charismat.Biocoenos.TopographSeasonal. Dis. select Vulnerable S-A relns. SubstrateProduct. Micro. Evol. EconomicTransitionsRepresent. H-A equil. ErosionPhenotypesFun.groupsDistinctive H-L equil. SpeciationFragmentsHeterog.AnomaliesTurbulence Macro. Evol Meta-popsEndemismExposureMixing Alt. S.Stats PatchinessUpwelling SymbiosesNutrientsDivergence Biomass Dis. Gases Ecol.Integ. AnoxiaErosion Desiccation GENETIC Structure Process SPECIES/ POPULATION Structure Process COMMUNITY Structure Process ECOSYSTEM Structure Process The components of marine biodiversity: Expanded from Zacharias and Roff 2000

Eco. level Approach ↓ Structur e ProcessStructureProcessStructureProcessStructureProcess Distinct. Habitats 1,2,3,4Inferred from structures 4,5,6A, 6B,7,8,9,10,11 1,3,4,5, 6,7 1,2,3,5, 6B,12, 13,16 Assumed or N/A 5,11,13, 18 1,2,3,4, 5,6,7,8, 9,10,15, 16,18 Repres. Habitats 1,2,3,4, 5 Inferred from structures 1,2,3,5,6A,6B,9,10, 11 5,6,7, 1,2,3,4,5, 6A,7,8,9, 10,11,13, 14,15,16 Assumed or N/A, 7,8 1,2,3,4,5, 6,7,8,9,10,12,14,15, 16,17 10,11,12, 13,14,18, 19 Fisheries Conserv. 1,2,3,4, 5 fish only Inferred from structures 1,2,3,6B,10 fish only 1,2,3,4, 5,6,7 fish only 6A,6B,9, 16 fish only Assumed or N/A N/A Coastal Zone Manag. N/A 2,3,4,5, 6,9,10, 12,14 2,3,4,5, 7,8,9,10,18 Sets of MPA’s 1,2,4,5Inferred from structures 1,2,3,5,6A,6B, 12 1,2,3,4, 5,6,7 6A,6B,7, 8,13,14 Assumed or N/A,1 1,2,3,4, 5 1,2,3,4, 5,6,7,8, 9,17 Networks of MPA’s 1,2,4,5Inferred from structures 3,11,12, 13 1,2,3,4, 5,6,7 10,14 Assumed or N/A N/A1,2,4,5, 7,17 How the elements of biodiversity are ‘captured’ by various conservation approaches GENETICECOSYSTEMCOMMUNITYPOPUL./ SPECIES

IT IS BETTER THAN THE BIASED TERRESTRIAL APPROACHES WE HAVE SO FAR! THIS APPROACH TO ‘CAPTURE’ VALUE MAY NOT BE PERFECT, BUT: