Washington tribal fishery resources

Potential vulnerabilities to ocean acidification

By Brad Warren, dir., Productive Oceans Partnership program

Brad.warren@sustainablefish.org

Some species of interest !  Salmon

!  Crab

!  Halibut & blackcod

!  Hake

!  Oysters, clams, other bivalves

!  Shrimp & prawns

!  Urchins, sea cucumbers, other echinoderms

Crosscutting issues 1. Physiology —not just calcification

!  Freshwater acidification and some lab marine studies show broad physiological impacts. Reduced:

!  Reproduction

!  Respiration

!  Growth

!  Hunting prowess

!  Predator avoidance

!  Immune response

!  Cardiac function

Crosscutting issues 2. Habitat

Corals Tipping point at pH 7.8:

below this level, “corals are completely

absent from these waters,” and

“community composition changes

dramatically.” (Hall-Spencer &

Rauer 2009)

Calcareous algae

Vent study: Covered 60% of seafloor

in areas at normal pH (8.1-8.2),

absent in areas below 7.9

(Hall-Spencer 2008) UPPER: Stylaster sp. with hagfish and shrimp. NOAA/OCNMS photo LOWER: Sebastes in a coldwater reef. UNEP photo.

Crosscutting issue 3. Foodweb impacts

!  Pteropods !  Copepods !  Foraminifera !  Euphausids

!  Planktonic fish larvae

UPPER: Pteropod (coexploration.org)

LOWER: Calanus pacificus from Dabob Bay (OSU)

Reliance on vulnerable species

Food web impacts

Salmon

!  Foodweb impacts: Heavy dependence on shelled prey, esp. in juvenile phases (pink, chum, sockeye): copepods, amphipods, pteropods, etc !  Pteropods: In aragonite-undersaturated water, visible

dissolution began within 48 hours. (Fabry, Orr, others)

!  Physiology: Salmon farmers concerned about possible reduced growth rate and other effects documented in acidified lakes, rivers.

Major feeding grounds of NW salmon

Does productivity = vulnerability?

!  West Coast upwelling zone —includes Puget Sound basin. !  Low pH, high CO2 documented during upwelling. !  Anthropogenic CO2 compounds effect. (Feely)

!  North Pacific !  NW salmon “turn right” to feed in north: cold, CO2-

enriched !  Saturation horizons at 100-200m !  Undersaturated conditions documented at surface

(Mathis).

What we know about the biological impacts of ocean acidification …on marine fish

Salmon Impacts

Pink salmon Juvenile diet can be 60% L. helicina pteropod (Armstrong et al 2005)

Results when pinks exposed to very low pH (mean 4.35)

!  Juveniles showed reduced growth.

!  Larvae showed 16.8x greater mortality rate than controls (47.1% compared to 2.8%) when exposed for 14 days. Most of this mortality was delayed, occurring within one week after removal from acidified water, with fish later kept at pH 7.5. Authors attributed mortality not to pH but to “aggregated dose” (i.e. lingering effects of acidosis, hypercapnia).

!  Females exposed to acid produced fewer oocytes with early yolk development.

!  Study used sulfuric acid, not CO2.

(Zelennikov et al 2006)

Salmon impacts in Nova Scotia

N.S. salmon associations seek 75% cut in U.S. “acid rain emissions” —SO2 & NOx from coal & oil.

Clues from Freshwater

Atlantic salmon !  “Acidification has caused the loss or reduction of

numerous Atlantic salmon populations on both sides of the North Atlantic.”

!  “Acid deposition peaked in the 1980s and resulted in both chronically and episodically acidified rivers.”

!  Acidification has affected Atlantic salmon in >50 rivers in Norway. Salmon are extinct in 18 of these rivers, catches reduced in the rest.

SOURCE: Kroglund et al 2008

Salmon: Clues from Freshwater

Reduced growth in fish & fish stocks

In acidified water, fish divert energy from growth and reproduction.

They do this to restore ion metabolism including acid-base balance.

“Taking into account the high sensitivity of fish eggs and larvae to acidification, these phenomena are probably a major contributory factor to the decline of fish stocks in acidified water.” (Wendelaer Bonga & Dederen, 2004)

Salmon: Freshwater clues

Other salmonids

!  Rainbow trout: reproduction affected below pH 5.5: oogenesis, spermatogenesis impacted. No eggs exposed to pH 4.5 survived to eyed stage. (Weiner et al, OSU 2004).

•  Much lower pH than expected in saltwater. To what extent

is freshwater experience applicable for predicting effects on marine fish?

Salmon: Clues from Freshwater

cordovatrout.blogspot.com

Acidification in lakes, rivers !  Lost and diminished populations.

!  Reduced growth.

!  Fish divert energy from growth and reproduction to restore ion metabolism including acid-base balance.

“Taking into account the high sensitivity of fish eggs and larvae to acidification, these phenomena are probably a major contributory factor to the decline of fish stocks in acidified water.” (Wendelaer Bonga & Dederen, 1986)

Salmon: Clues from Freshwater

Aluminum toxicity from acidified rivers

!  Aluminum and other metals are “mobilized” in acidified rivers; aluminum a major source of mortality. Vulnerability high in smolts, but less in parr. (Kroglund et al 2008)

!  “Massive salmon mortality” at Norwegian salmon farms: “probable cause” was aluminum concentrating in gills, due to storm runoff from acidified, aluminum-rich rivers. (Bjerknes et al 2003)

Salmon: Clues from Freshwater

Crab • Strong ion regulation found in some species (Ries et al and Meizner et al 2009, but will it help in wild?

• Unknown vulnerability at larval, moult stages.

•  Some evidence for increased shell growth to deal with high CO2 but likely at cost (Ries et al 2009).

• Cost of compensatory shell growth? Brittle stars increase shell growth but suffer muscle wastage (Wood et al 2008).

Crab – mixed effects !  Dungeness crab showed high tolerance for CO2 (down to pH 7): better able to compensate

for acidosis than deepwater Tanner crab by accumulating bicarbonate to elevate haemolymph pH. Authors hypothesized that deep-sea animals are less able to cope than shallow dwellers, which live in a more variable pH environment. (Pane & Barry 2007).

!  Ocean acidification effects on larval Dungeness crabs being studied WWU’s Shannon Point lab. (WWU press release 2009).

!  Velvet swimming crab N puber: Hardy. Mostly bicarbonate compensates for acidosis. Partly supplied by dissolving exoskeleton. BUT after 16 days, extracellular acidosis at pH 6.74; marked acidosis after 24 hours at pH 6.05. Locally acute conditions “likely to compromise even this species.” (Spicer et al 2006)

!  However, other crabs exposed to low pH showed reduced metabolic activity, regardless of their habitat (Spicer 2007).

!  Spider crab (H araneus): Tolerance for warm conditions declined at moderately elevated pCO2. From 25C at present normal CO2 level (380 ppm), critical temperature dropped to 23.5C at 710 ppm . Result: oxygen & capacity limitation in this eurythermal coldwater crab. (Walther et al 2009).

!  Thermal tolerance might be an important compounding factor for crustaceans, e.g. Southern New England lobster recruitment collapse. (ASMFC 2010)

Mechanisms of resilience in some crustaceans?

Resilience to high CO2 “has only been observed in adults/juveniules of active, high metabolic species with a powerful ion regulatory apparatus. However, while some of these taxa are adapted to cope with elevated pCO2 during their regular embryonic development, unicellular gametes, which lack specialized ion-regulatory epithelia, may be the true bottleneck for ecological success—even of the more tolerant taxa.” (Meizner et al 2009)

Crabs in Hood Canal

Halibut & blackcod – some resilience?

Halibut spawn in deep water (180-450 m) along slope. Eggs develop at depth, are planktonic up to 7 mos. Larvae move to shallows (2-50 m), become largely piscivorous at about 30 cm.

Blackcod spawn at 300-700 m. Eggs & larvae remain in deep water. Young of year (~8 cm) eat zooplankton in first weeks.

Unknown: Does tolerance of high CO2 in early life stages indicate resilience to increases in that level? Or are they near physiological limits

Hake & other gadoids !  Roughly similar to pollock, cod, so research on these

species may provide clues to impacts.

!  Cod: sperm motility unaffected by OA. (Frommel et al 2010)

!  NOAA’s Manchester Research Station recently completed life-cycle of P-cod in capacivity, an enabling step for future studies on OA and climate impacts.

!  Pollock: preliminary evidence of bicarbonate increase indicating buffering. Cost?

!  Pollock study pending at UAF under Jeremy Mathis looking at age 0, age 1: Hatching success, % mortality, growth rate, body condition, metabolic enzyme activity in age-1 juveniles, stress hormone levels.

Oysters, clams, other bivalves

High dependence on aragonite. Crassostrea gigas after 48 hours incubation. Frame a is the control. It was incubated in standard seawater (8.2 pH). Frames c, e, and g were incubated in pH 7.4 seawater (Kurihara, 2007).

Northwest oyster seed crisis Pacific oyster larvae fail

70-80% loss of production in 2007-2008 at both major hatcheries. At Whiskey Creek, oyster larvae dissolved, vanished in tanks. Even hard-fouling of intake pipes ceased.

Little or no commercial-scale wild “set” of oysters in Willapa Bay since 2005.

Pacific oyster larvae growing at Taylor hatchery on Dabob Bay. Billions of these were lost.

T= 0 hours T = 24 hours T = 72 hours

SEM’s of larval-stage M. mercenaria reared in aragonite-undersaturated seawater. Size ! 100µm, mag. = 370-400X, pH = 7.5, Ωaragonite = 0.5.

Dissolving clams

Reduced growth & calcification !  Bay scallop A. irradians: up to 70% mortality in larvae, delayed settlement (up to 5 weeks).

(Talmage & Gobler 2009)

!  Greenlip abalone: H. laevigata 5% & 50% reduced growth at pH 7.78 and 7.39 (Harris et al 1999)

!  Blacklip abalone: H. rubra 5% & 50% reduced growth at pH 7.93, 7.37 (Harris et al 1999)

!  Pacific oyster C. gigas: at pH 7.4, development to D-veliger stage was 7.3% of normal (only 5%, compared to 68% normal). (Kurihara et al 2007)

!  Eastern oyster C. virginica (Miller et al 2009)

!  Hardshell clam M. mercenaria (Green et al 2004, 2009)

!  Soft-shell clam M. arenaria (Salisbury et al 2008)

!  Mediterranean mussel M. galloprovincialis (Kurihara et al 2008)

!  Sydney rock oyster S. glomerata (Watson et al 2009)

!  Blue mussel M. edulis: thinner shells (D-veliger), hatching declined 24% at pH 7.6 (Gattuso et al 2010). At 740 pC02, 25% reduced calcification (Gazeau et al 2007)

Bivalves under ocean acidification

Tatoosh Island

Resistant bivalves? No reduction in growth, calcification •  Suminoe oyster C ariakensis (Miller et al 2009)

Some resistance? •  Bivalves with periostricum?

•  Possibly Olympia oyster? But 41% less growth at 970 ppm after 19 days. (NSF 2010)

Shrimp !  Impacts on shrimp still

poorly known, but preliminary evidence points to negative effects.

!  Shrimp farmers now avoid mangrove swamps, partly because “acidic soil found in mangrove areas is unfavorable for shrimp farming.” (Gillett 2008)

Shrimp (Palaemon pacificus)

an intertidal shrimp (possibly accustomed to some variability in pH)

55-65% die at 1,000-1900 ppm CO2

Growth, moulting, egg production reduced.

Antenna are stunted.

!  2100: 1,000 ppm. 55% die (normal (Control mortality = 10%) at 30 weeks.

!  2300: 1,900 ppm. 65% die (normal (Control mortality = 5%) at 15 weeks.

(Kurihara et al, 2008)

Krill

!  At CO2 levels projected to occur worldwide 300 years in future (already found in some waters), krill eggs failed to hatch, larvae developed irregular form, became less active. (Nicol 2008)

!  Antarctic krill mortality rises with exposure time at pH 7.6 (a level already found seasonally in NW seas). (Yamada and Ikeda 1999)

Urchins, cukes, other echinoderms

!  Endoskeletons often use magnesium

calcite form —soluble.

!  Urchin recruitment correlated to downwelling: link

to high CO2? (Steele 2010). Fertilization declines with rising CO2 (Kurihara and Shirayama 2004)

!  Deformations, reduced growth, fitness in urchin larvae.

“With increased acidity/pCO2 larval size decreased and there was an increase in abnormal development” in Priest-hat urchin. (Sheppard Brennand et al 2010).

NWIFC

Hatfield Marine Sci Center— coast-nopp.org

Cliff Cultee, Lummi Nation fisherman

Red urchin

Global catch of D gigas > 600,000 t (FAO)

Squid Minimal armhook squid B. anonychus: a key prey for juvenile pink, sockeye salmon during diet shift. Abundance is a control on adult salmon biomass. High metabolic rates may lead to high sensitivity to CO2 (Seibel 2007).

Humboldt squid (possibly fastest growing fishery In the world) D. gigas: metabolic depression under combined high CO2 (1,000 ppm) and warming projected by end of 21st Century (25C) Max. MR declined 31%. (Rosa & Seibel 2008).

LEFT: Humboldt Squid caught off La Push in 2004

Gastropods Horned turban turbo cornutus

sold in sushi restaurants as Sazae Growth and shell extension reduced at 560 ppm (Shirayama & Thornton 2005).

Snail shells dissolve at pH 7.6 near CO2 vent (Hall-Spencer 2008)

Pinto abalone Once abundant in Puget Sound, now only a few thousand remain.

Bibliography Salmon in acidified Rivers Bjerknes et al 2003: Aluminum in acidic river water causes mortality of farmed Atlantic Salmon

(salmo salar L.) in Norwegian fjords, Marine Chemistry, Vol 83, iss. 3-4, Nov 2003. Kroglund et al 2008: Water quality limits for Atlantic salmon (Salmo salar L.) exposed to short

term reductions in pH and increased aluminum simulating episodes, Hydrology and Earth System Sciences, 12, 491-507, 2008.

Wendelaer Bonga & Dederen, 1986: Effects of acidified water on fish, Endeavor, Vol 10, Issue 4, 1986, p 198-202

Acid Rain Kills Nova Scotia Rivers, Atlantic Salmon Federation flyer, online at www.asf.ca/docs/issues/acidrain.pdf

Foodweb & Physiology

Aydin et al 2005: Linking oceanic food webs to coastal production and growth rates of Pacific salmon (onchorynchus spp.), using models on three scales. Deep Sea Research II 52 (2005) 757-780.

Frommel et al 2010: Effect of ocean acidification on marine fish sperm (Baltic cod: Gadus morhua), Biogeosciences Discuss., 7, 5859-5872, 2010. www.biogeosciences-discuss.net/7/5859/2010

Kurihara et al 2004: Sub-lethal effects of elevated concentration of CO2 on planktonic copepods and sea urchins. J. Oceanogr., 60, 743-750, 2004.

Zelellikov et al 2007, Effect of water acidification on the oogenesis of the pink salmon Oncorhynchus gorbuscha, Journal of Ichthyology, Vol 47, No 3, Aril 2007, p 254-257.

Armstrong 2005: Distribution, size, and interannual, seasonal and diel food habits of northern Gulf of Alaska juvenile pink salmon, Oncorhynchus gorbuscha, Deep Sea Research II, 52: 247-265.

Habitat

Hall-Spencer & Rauer 2009: Champagne Seas—Foretelling the Ocean’s Future? Current: The Journal of Marine Education, Vol 25, No. 1, 2009

Hall-Spencer et al: 2008:Volcanic carbon dioxide vents show ecosystem effects of ocean acidification, Nature 454, 96-99, 3 July 2008.

Bibliography cont’d Bivalves

NSF 2010 Releas describing findings of NSF-funded research by Brian Gaylord and colleaguees at Bodega Marine Lab, UC-Davis. Online at http://www.nsf.gov/news/news_summ.jsp?cntn_id=116767

Harris et al 1999: Effect of pH on growth rate, oxygen consumption rate, and histopathology of gill and kidney tissue for juvenile greenlip abalone, Halitosis laevigata and blacklip abalone, Halitosis rubra leach. Journal of Shellfish Research, 18: 611-619.

Kurihara et al 2007: Seawater carbonate chemistry and processes during experiments with Crassostrea gigas, 2007, doi:10.1594/PANGAEA.721193. Also referencing Kurihara et al 2007, Effects of increased seawater pCO2 on early development of the oyster Crassostrea gigas, Aquatic Biology, 1(1), 91-96.

Hall-Spencer et al: 2008:Volcanic carbon dioxide vents show ecosystem effects of ocean acidification, Nature 454, 96-99, 3 July 2008.

Urchins, Echinoderms, Gastropods

Sheppard Brennand et al 2010: Ocean acidification alters skeletogenesis and gene expression in larval sea urchins, Mar Ecol Prog Ser 398: 157-171, 2010, online at http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0011372

Steele 2010: presentation at workshop “Ocean Acidification Effects on the West Coast Shellfish Industry,” Costa Mesa CA, July 7, 2010.

Shirayama & Thornton 2005: Effects of increased atmospheric CO2 on shallow water marine benthos, Journal of Geophysical Research, Vol 110, C09S08, 7 pp, 2005.

Bibliography, cont’d Crab

ASMFC 2010: Recruitment Failure in the Southern New England Lobster Stock, American Lobster Technical Committee, Atlantic States marine Fisheries Comission, http://www.asmfc.org/ (under “Breaking News” tab). April 2010 Meizner et al 2009: Physiological basis for high CO2 tolerance in marine etcothermic animals: pre-adaptation through lifestyle and ontogeny? Biogeosciences Discussions, 6, 4693-4738, 2009. www.biogeosciences-discuss.net/6/4693/2009/ Pane & Barry 2007: Extracelluar acid-base regulation during short-term hypercapniais effective in shallow-water crab but ineffective in a deep-sea crab, Marine Ecology Progress Series, Vol 334:1-9, 2007. Spicer et al 2006: Influence of CO2-related seawater acidification on extracellular acid-base balance in the swimming velvet crab Necora puber, Mar Biol (2007) 151:1117-1125 (published online Dec 2006). Walther et al 2009: Impact of anthropogenic ocean acidification on thermal tolerance of the spider crab Hyas araneus, Biogeosciences, 6, 2207-2215, 2009. www.biogeosciences.net/6/2207/2009/ WWU press release 2009: Shannon Point Scientists Receive $557,000 Grant to Study Ocean Acidification, www.piersystem.com/go/doc/1538/477143/

Gillett 2008: A global study of shrimp fisheries, FAO Technical Paper 475 Nicol 2008: Krill face shell shock, Unitas, University of Tasmania, October 2008 issue 324 Yamada & Ikeda 1999: Acute toxicity of lowered pH to some oceanic zooplankton. Plankton Biology and Ecology, 46: 62-67.

Shrimp & Krill

Bibliography, cont’d

Squid

Rosa & Seibel 2008: Synergistic effects of climate-related variables suggest future physiological impairment in a top oceanic predator. PNAS Vol 105, No 52, 20776-20780, Dec 30, 2008.

Seibel 2007: On the depth and scale of metabolic rate variation: scaling of oxygen consumption and enzymatic activity in the Class Cephalapoda (Mollusca). Journal of Experimental Biology, 210: 1-11