what is PISCO The Partnership for Interdisciplinary Studies of Coastal Oceans is a long- term program of scientific research and training dedicated to advancing the understanding of the California Current Large Marine Ecosystem along the U.S. West Coast. PISCO is pioneering an integrated approach to studying this complex, rich, and economically important environment. PISCO is distinguished by its interdisciplinary approach, large geographic extent, and decades-long time frame. PISCO conducts monitoring and experiments along more than 1,200 miles (2,000 kilometers) of coastline, as well as laboratory and theoretical studies. The research incorporates oceanography, ecology, chemistry, physiology, molecular biology, genetics, and mathematical modeling to gain novel insights into systems ranging from individual animals and plants to the whole ecosystem. PISCO’s findings apply to conservation and resource management issues. PISCO scientists participate in local, regional, national, and international initiatives for marine environmental planning. Through its university courses, PISCO helps to train the next generation of scientists in interdisciplinary approaches to marine research and policy. Established in 1999 with funding from The David and Lucile Packard Foundation, PISCO is led by scientists from Oregon State University (OSU), Stanford University’s Hopkins Marine Station, University of California at Santa Cruz (UCSC), and University of California at Santa Barbara (UCSB). As of 2005, core PISCO activities are funded by collaborative grants from The David and Lucile Packard Foundation and the Gordon and Betty Moore Foundation. The core support and additional funding from diverse public and private sources make this unique partnership possible. PISCO View from the Wave Crest Coastal Connections Volume 5

Table of Contents ISCO Coastal Connections is an annual publication of the Partnership for 1 View from the Wave Crest Interdisciplinary Studies of Coastal Oceans 2 Patterns of Change P (PISCO). We welcome you to the fifth issue in the Reproductive Hotspots series that highlights major findings, research projects, Fish Population Genetics as well as outreach and education. Physics of Rocky Shores The interdisciplinary PISCO consortium is an interconnected group of estab- Identifying Fish Birthplaces lished scientists and postdoctoral fellows, science and policy coordinators, data managers, graduate students, and research technicians. The consortium’s work 6 Oceanographic Frontiers is further strengthened by its collaborations with government agencies, non- Monitoring Oceanography government organizations, and academic institutions. This issue of PISCO Modeling Drifting Young Coastal Connections reflects these cumulative efforts and their value for marine Currents and Dispersal Paths policy and management. Articles in this year’s “Patterns of Change” section feature some of our latest 10 Ecological Linkages findings about variations over time and distance in the marine ecology and Changes in Kelp Forests oceanography of the U.S. West Coast. “Oceanographic Frontiers” describes Oceans Affect Shore Ecology PISCO’s extensive ocean-observing network and our research into oceano- Effects of Climate Warming graphic processes in coastal waters. The “Ecological Linkages” section explores new findings about the connections between oceanography and ecological 14 Interdisciplinary Training changes. Results from PISCO’s graduate student programs, including our new & Research marine policy course, are highlighted in “Interdisciplinary Training.” “Sharing PISCO Training Courses the Science” showcases examples of PISCO’s initiatives to communicate scien- 1 Student Showcase tific findings and methods to broader audiences. We invite you to enjoy this issue of PISCO Coastal Connections and the 16 Sharing the Science achievements described on the following pages. Teaching Groups to Monitor PISCO Advises California Straight Talks on Fishing Methods of Stock Assessment

PISCO Coastal Connections Program Coordinator: Kristen Milligan PISCO Coastal Connections Coordinators: Satie Airamé, Liz Riley, Cinamon Vann, Amy Windrope Editor & Writer: Peter H. Taylor Creative Director: Monica Pessino Graphics Assistant: Julia Kwinto GIS Support: Will McClintock Line Drawings: Linda D. Nelson Cover photo: Painted greenling (Oxylebius pictus) © 2006 Luke Miller. Cover photo insets, top to bottom: Wyatt Patry, Francis Chan, Jane Lubchenco, Monica Pessino. Opposite page photos, left to right: Gretchen Hofmann, Giacomo Bernardi, Luke Miller, Amy Wagner PISCO Coastal Connections is a publication of PISCO principal investigators (top to bottom, left to right): Libe Washburn (UCSB), Pete Raimondi (UCSC), Steve the Partnership for Interdisciplinary Studies of Gaines (UCSB), Mark Denny (Stanford), Margaret McManus (UCSC), Jane Lubchenco (OSU), Jack Barth (OSU), Robert Coastal Oceans (PISCO). Contents © 2006. Warner (UCSB), Steve Palumbi (Stanford), George Somero (Stanford), Mark Carr (UCSC), and Bruce Menge (OSU). For more information about PISCO or to join Not shown: Gretchen Hofmann (UCSB). Photo: Satie Airamé the mailing list for future publications, please contact the consortium at the addresses listed on the back cover.

0)3#/#OASTAL#ONNECTIONS„6OLUME patterns improve management of marine ecosystems (see article, next page). are likely to have reproductive hotspots, and identifying these hotspots could barnacles from Cape Foulweather (see figure above). Other marine species differences, as Cape Perpetua barnacles produced 120 times more young than major foods. In targeted experiments, the scientists uncovered even greater Cape Perpetua contain much more phytoplankton, one of the barnacles’ difference may arise from differences in food availability. The waters near many offspring as barnacles near Cape Foulweather, Oregon. This reproductive coast near Cape Perpetua, Oregon, produced approximately five times as In monitoring surveys, PISCO scientists Geographic DifferencesinBarnacleReproduction found that barnacles living on the

OFchange Hotspots for Reproduction and Conservation

arine scientists have long suspected that populations of invertebrates living in different places M in the sea may differ in their production of young, but scientists have rarely tested this idea empirically. Using the acorn barnacle (Balanus glandula) as a model, PISCO/OSU doctoral student Heather Leslie Barnacles (Balanus glandula) on the rocky shore. examined geographic variation in barnacle reproduction along the Oregon Photo: Sheri Etchemendy coast to find out whether food supply affected the number of offspring produced. Her research builds on PISCO’s prior findings that phytoplankton, which barnacles eat, are abundant near Cape Perpetua but scarce near Cape Foulweather. Leslie’s research offers an example of how marine scientists can effectively gather information on spatial variation in reproduction and Science of Marine Reserves link it to ecological and oceanographic processes. Leslie collaborated on the Marine reserves offer greater protec- research with PISCO principal investigators Jane Lubchenco and Bruce Menge, tion than any other type of marine postdoctoral fellow Francis Chan, and OSU honors student Erin Breck. protected area (MPA) by completely The research highlights the value of understanding how ecological and protecting animals, plants, and their oceanographic processes influence marine populations. Apparently similar habitat from removal or alteration. habitats along the coast differ markedly in their ecological functioning (see Other MPA designations may allow findings, opposite page). Particular sites may be especially significant sources certain human uses, like recreational of young for some marine invertebrates, serving as reproductive hotspots. 3 fishing. Design of marine reserves and Leslie’s findings are relevant to design of marine protected areas because they other protected areas can be informed suggest the potential to identify and conserve sites with high reproductive by scientific research regarding species rates for certain species. distributions, abundance, dispersal, population replenishment, and inter- Heather Leslie is now a postdoctoral fellow at Princeton University. actions with other species. Research Publication: Barnacle reproductive hotspots linked to nearshore ocean conditions. Proceedings of shows that marine reserves can help the National Academy of Sciences 102 (2005): 10534–10539. to protect marine habitats and species, and in some cases restore populations of depleted species. Fish, shellfish, and seaweeds inside marine reserves tend to be larger, more abundant, and more diverse than they are in non-reserve areas. These benefits of marine reserves can boost marine ecosystem resilience and productivity. For more information about the science of marine reserves, visit www.piscoweb.org.

Heather Leslie studying barnacle reproduction at Fogarty Creek, along the Oregon coast. Photo: Jane Lubchenco

0)3#/#OASTAL#ONNECTIONS„6OLUME Genetics Show Distinct Rockfish Populations PISCO/UCSC graduate student Martha Burford is studying geographic varia- tion in the population genetics of blue rockfish (Sebastes mystinus) and kelp rockfish (S. atrovirens) along the California coast. Adult rockfishes generally do not travel far from the kelp beds and rocky reefs where they live. However, long-distance dispersal of their young might link the state’s rockfishes into one genetically similar population. Burford uses genetic markers, called microsat- ellite loci, to detect genetic differences among adults and juveniles of both species. Her research reveals substantial geographic variation in the population genetics of juvenile rockfishes in California’s waters, indicating that the state has multiple genetically distinct populations. Burford’s findings have implica- tions for fisheries management. For example, a network of several marine pro- tected areas or marine reserves might protect the genetic diversity of rockfish better than one large area.

Caption Martha Burford prepares DNA from blue rockfish for analysis. Photo: Lydia Bergen Finding Shelter on the Rocks PISCO/Stanford graduate student Michael “Moose” O’Donnell explored the fluid mechanics of wave-battered shores to understand how topography affects ecological patterns. His research revealed that crevices on rocky shores do not necessarily protect invertebrates and seaweeds from pounding waves—and may, in fact, do the opposite. Depending on size and orientation, a crevice can funnel water, creating intense wave forces. Yet, in some cases, snails, 4 seaweeds, and other shore-dwelling species actually benefit from the stronger wave forces, as water splashing from the crevices enables them to live higher on the rocks than they otherwise could. In contrast, O’Donnell has found that mussel beds do create shelter for small animals. Wave forces experienced by a snail, for example, drop by 40 percent within 10 centimeters of a mussel bed—and by 90 percent within one centimeter. This research provides insight into how components of the physical environment interact to create habitable space for marine animals and plants along the coast.

Michael O’Donnell is now a postdoctoral fellow at the University of California, O’Donnell measures wave forces in mussel beds. Santa Barbara. Photo: Luke Miller

crevice

Green box indicates location of close-up photo (left). Photo: Michael O’Donnell O’Donnell has found that rocky crevices can increase the wave forces experienced by marine animals and seaweeds. Heights of bars indicate wave forces measured by wave-force recorders (white balls below bars). When a two-meter wave hits, the force in a small crevice (red bars) can be double that outside the crevice (orange and green bars).

0ARTNERSHIPFOR)NTERDISCIPLINARY3TUDIESOF#OASTAL/CEANS patterns of change 5 - Limnology and They measured seven seven measured They

). These signatures ). These signatures 0)3#/#OASTAL#ONNECTIONS„6OLUME Sebastes atrovirens 50(2005): 1529–1542. (see figures and caption). (see figures Results suggest that otoliths could be used to identify the birthplace of adult along and temporal patterns in otolith signatures after geographic rockfishes, the connections Ultimately this method could reveal understood. the coast are among fish populations, enabling better fisheries management. ResearchersWarner are Jennifer (PISCO/UCSB); Caselle, in the otoliths of an open-coast fish. Publication: Natal trace-elemental signatures GeorgesOceanography and Stephen Paradis, Swearer Michael (University Sheehy, and of Melbourne,Robert Australia). Natural Tags Indicate Fish Birthplaces Fish Indicate Tags Natural that water chemistry along the open PISCO/UCSB scientists have discovered the otoliths, or in chemical signatures coast varies enough to leave identifiable ( ear bones, of newborn kelp rockfishes of rock to be useful as natural “tags” that identify the birthplaces may prove fishes caught as adults. along the mainland and areas three from The scientists collected larval rockfish the tiny fish otoliths from islands near Santa Barbara. The scientists removed (see sidebar). and analyzed the chemical composition using mass spectrometry the They found detectable levels of several trace elements in the otoliths, and in a way that was distinctive among the study sites varied chemical signatures - - - - - Trace Trace

PISCO Coastal PISCO Photo: Marc Chamberlain Photo: ). ). to 4. 1

The otolith grows by add by The otolith grows

, Volumes , Sebastes atrovirens Kelp rockfish ( rockfish Kelp tion about the technique, see technique, the about tion Connections and where it traveled. For more informa more For it traveled. and where tists intend to use the otolith like a natural tists intend to use the otolith like a fish was born of where “flight recorder” layers. By matching the layers’ signatures to signatures By matching the layers’ layers. the scien particular places in the ocean, eters to measure the trace elements in the eters to measure are recorded in the otolith layers. PISCO in the otolith layers. recorded are scientists use lasers and mass spectrom so as the fish travels, these differences these differences so as the fish travels, ing calcium layers, like rings in a tree. like ing calcium layers, elements vary among places in the ocean, ocean water into a fish’s ear bone, or oto ear bone, ocean water into a fish’s as it grows. lith, Trace elements are incorporated from from incorporated elements are Trace elements: strontium (Sr), barium (Ba), lead (Pb), magnesium (Mg), manganese (Mn), iron (Fe), and zinc (Zn). Because the relative amounts of the elements varied from site to site, the site to site, amounts of the elements varied from Because the relative and zinc (Zn). (Fe), iron manganese (Mn), (Mg), magnesium lead (Pb), barium (Ba), (Sr), strontium elements: each fish was born. tags identifying where trace elements acted as distinctive Deciphering the Tags PISCO/UCSB scientists collected larval kelp rockfishes from three areas near Santa Barbara and analyzed the trace elements in the fishes’ otoliths, or ear bones. the trace elements in the fishes’ otoliths, near Santa Barbara and analyzed areas three from PISCO/UCSB scientists collected larval rockfishes kelp Network of Oceanographic Moorings Locator Map

NMS = National Marine Sanctuary SBC-LTER = Santa Barbara Coastal Long-Term Ecological Research

PISCO scientists are working to identify how oceanographic conditions help shape coastal ecosystems. To study the marine environment near shore, PISCO deploys and maintains dozens of oceanographic moorings from Oregon to southern California. Partnerships with the National Oceanic and Atmospheric Administration’s (NOAA’s) National Marine Sanctuaries (NMS) program and the National Science Foundation’s Santa Barbara Coastal Long-Term Ecological Research Program (SBC-LTER) enable greater coverage and expertise. PISCO’s database experts and scientists are developing new systems to manage the vast amounts of oceanographic data from PISCO moorings. The data are available by contacting PISCO staff, or through online data catalogs at www.piscoweb.org. oceanographic frontiers Monitoring Detects Ocean Anomalies

Oregon n early summer 2005, many people began to notice unusual changes along the North American west coast—warmer ocean temperatures I near the shore, decreased concentrations of plankton in some coastal waters, a drop in groundfish catches, and an increase in dead seabirds found on beaches. A delay of nearly two months in the onset of normal springtime, southward Central California winds resulted in little or no upwelling of cold, nutrient-rich deep waters. This led to warm surface waters, scarce nutrients, and lowered productivity along the coast. During this apparently anomalous event, PISCO researchers detected unusual ocean conditions and consequent ecological impacts. • Warmer water (figure, left) – Most pronounced in Oregon during the late spring, the water reached 18 degrees Celsius—5 to 7 degrees above normal. Coastal waters of California were 1 to 3 degrees Celsius warmer Southern California than normal early in 2005. • Changes in phytoplankton – Surveys in Oregon found approximately half as much phytoplankton in May and June as in previous years. In Mon- terey Bay, a shift occurred in the phytoplankton; dinoflagellates dominated in summer 2005, whereas diatoms were dominant in 2002 (Jim Sullivan, University of Rhode Island). • Low population replenishment and larval abundance – Mussel 7 population replenishment was the lowest ever observed in Oregon for at least the last decade. Along the central coast of California, six species of Red lines on the graphs indicate average daily water tem- rockfish failed to add young to their populations in kelp forests. Offshore peratures on the ocean surface in 2005 at PISCO moor- ings. Blue lines show the averages for 2001 to 2004 and surveys by NOAA revealed very few larvae. variation around this average. When the red line is above or below the blue region, temperatures in 2005 differed PISCO scientists are now working with researchers at other institutions to significantly from previous years. Moorings in Oregon are evaluate the ecological impacts of the unusual period of warm water. deployed only from spring to fall each year because of intense winter storms; therefore data are not available for PISCO researchers are Jack Barth, Francis Chan, Anthony Kirincich, Jane winter and early spring months. Lubchenco, Bruce Menge (OSU); Mark Carr, Patrick Drake, Margaret McManus, Pete Raimondi (UCSC); Jennifer Caselle, Chris Gotschalk, Libe Washburn (UCSB).

The Cause: Delay in Southward Winds Unusual wind patterns were the driving force behind major ecosystem anomalies along the Oregon coast in 2005. PISCO/OSU researcher Jack Barth and OSU researchers Steven Pierce and Renato Castelao analyzed wind records from the last 20 years for compari- son with 2005. Southward winds drive the upwelling of cold, nutrient-rich waters to the ocean surface. One measure used by researchers to determine the onset and magnitude of upwelling is called “cumulative wind stress.” The figure (right) shows cumulative south- ward wind stress (negative values) as measured from an offshore NOAA buoy. Normally, these winds blow strongly enough for upwelling to fuel the coastal food web through the summer. In 2005, upwelling was delayed by about two months due to lack of favorable winds. From June to July 2005, cumulative wind stress was the least ever observed in the past 20 years. In late July 2005, upwelling-favorable wind stress finally reached more typical conditions, but then it went into overdrive with more persistent upwelling than normal. By mid-September, the total amount of upwelling caught up to the historical average—but it was too late for the fish, birds, and other species that rely on upwelling during summer. The graph (right) shows the 20-year average (black line) for cumulative wind stress and the variation around this average (shaded); the blue line indicates conditions in 2005.

0)3#/#OASTAL#ONNECTIONS„6OLUME Modeling Dispersal of Young Fish Off southern and central California, oil and gas platforms serve as reef-like homes for large numbers of juvenile rockfishes. The juveniles settle at the platforms or at natural reefs after drifting with ocean currents. Scientists, resource managers, and conservationists have debated the ecological consequences for rockfishes if decommissioned platforms are removed. Do the platforms help boost rockfish numbers by providing additional habitat, or do the platforms simply attract fish that otherwise would settle at natural reefs? In collaboration with scientists funded by the U.S. Minerals Management Service, PISCO/ UCSB scientists developed a model that offers insights. Using data from high-fre- quency radar that measures the speed and direction of ocean currents, the scientists simulated the likely paths of juvenile rock- fishes drifting near a platform located off Lines indicate dispersal trajectories of barnacle young in the oceanographic model. Stars indicate points on the shore Point Conception. The model showed that where young were released in the models. During upwelling conditions, young tended to be carried southward and away from shore (black lines). During periods without upwelling, called “relaxation,” more young were retained and most of the juveniles would drift offshore, moved to the shoreline (red lines). away from suitable natural reefs, if there were no platform at which they could Models Link Oceanography and Animal Behavior settle. This result suggests that oil and gas Because many fish and invertebrate species release their young into the water platforms provide supplemental habitat for 8 to spend weeks or months drifting, the dispersal of young serves as an im- juvenile rockfishes that otherwise would portant link among populations. A major goal of PISCO is to understand the perish offshore. However, the model does interconnections of fish and invertebrate populations. Tracking the tiny young is not yet account for swimming efforts extremely difficult, so PISCO is developing numerical models, based on oceano- since little is known about the behavior graphic and ecological data, to simulate the dispersal. Such models traditionally of juvenile rockfish in the coastal ocean. treat the young as passive, drifting particles. Yet the young of many species PISCO scientists now seek to incorporate actively regulate their depth in the water, moving upward or downward over behavior into the model, enabling more ac- time. Currents move in different directions and speeds at different depths, so curate understanding of the role of oil and this behavior can affect the path of dispersal. gas platforms as rockfish habitat. PISCO is working to integrate the behavior of young invertebrates and fish into Researchers are Brian Emery, Milton Love, dispersal models to better mirror actual conditions. Scientists at PISCO/UCSC Mary Nishimoto (UCSB), Libe Washburn have created a model for Monterey Bay that combines physical oceanographic (PISCO/UCSB), and Carter Ohlmann (Scripps). data with information on barnacle larval growth and behavior. The coupled Publication: Do oil and gas platforms off Cali- physical-biological model simulates current speeds and directions throughout fornia reduce recruitment of bocaccio (Sebastes paucispinis) to natural habitat? An analysis based the bay at various depths. During the simulation, the model releases barnacle on trajectories derived from high-frequency radar. larvae at points along the coast and then tracks them as they drift. The larvae in Fisheries Bulletin, in press. the model change depth with each developmental stage, just as real barnacle larvae do. Taking into account each larva’s depth over time and the currents at that depth, the model determines the trajectories of the larvae through the ocean. Larvae may be swept out to sea, where they would perish, or they may be retained near the coast, where they could settle and mature. The scientists found that the proportion of larvae retained—and therefore potentially surviv- ing to adulthood—depends on where the larvae were born and the oceano- graphic conditions during the period that they drift (see figure above). Researchers are Anna Pfeiffer-Hoyt, Margaret McManus (PISCO/UCSC/ University of Hawaii), Pete Raimondi (PISCO/UCSC), and Yi Chao (Jet Propulsion Laboratory). Publication: Dispersal of barnacle larvae along the central California coast: A modeling study. Young bocaccio rockfish (Sebastes paucispinis). Limnology and Oceanography (in review). Photo: Donna Schroeder

0ARTNERSHIPFOR)NTERDISCIPLINARY3TUDIESOF#OASTAL/CEANS oceanographic frontiers 9 - - - 0)3#/#OASTAL#ONNECTIONS„6OLUME 50(5), 2005, 1473–1479. Limnology and Oceanography - biology their affected settlement patterns. barnacles (upper graph) barnacles (upper graph) (lower and mussels settled along graph) eastern the island’s while lines), (red shores settled along the few (blue shores western suggesting that the lines), carried warmer current to the young more mus In addition, island. a long sels settled over winter period between whereas and summer, barnacles settled in pulses during the spring indicating and summer, in that differences Because of ocean currents, the western shores shores the western Because of ocean currents, colder have California, of Santa Cruz Island, water (blue colors) than the eastern shores, warmer waters (yellow-red bathed in which are colors). of young Large numbers cally and tended to be simultaneous around the island. Mussels, in contrast, the island. Mussels, around cally and tended to be simultaneous in the number markedly continually over time, and sites differed settled more high numbers brought the warm currents of young. The findings suggest that few larvae to western brought of larvae to eastern sites, while cold currents and barnacle behavior or development appar in mussel sites, and differences The findings may of population replenishment. patterns ently caused different such as areas of marine protected effects have implications for the design and established in the Channel Islands. those recently Publication: Recruitment of intertidal invertebrates and oceanographic variability at Santa Cruz Island, California, U.S.A. Bernardo Broitman used the unique setting to examine how currents affect affect currents to examine how the unique setting used Broitman Bernardo the rocky arriving to inhabit of young invertebrates and abundance the timing replenishment the population the scientists monitored For five years, shore. every at several sites on the island; of mussels and barnacles two to three collectors fastened to the young that had settled onto months, they counted sensing remote used satellite Over the same period, the researchers the rocks. island. the around to track ocean currents of seawater temperatures differed and rates of population replenishment Oceanographic conditions and western sides. Persistently warm eastern the island’s strikingly between barnacles and numbers of young large where eastern shore, water bathed the at western sites, and few bar cold water prevailed mussels settled. Meanwhile, in the settlement patterns also emerged settled. Differences nacles and mussels of barnacles happened sporadi Population replenishment of the two species. Currents Influence Population Replenishment Population Influence Currents convergence California, lies at the Santa Barbara, of Santa Cruz near The island and Blanchette scientists Carol PISCO/UCSB ocean currents. of two major - - -

will enable better resource management. will enable better resource to understanding fluctuations in abundance and of shellfish and other invertebrates local populations. The research is essential The research local populations. complex effects of ocean circulation on of ocean circulation complex effects barnacles to replenish of young the arrival ecological monitoring is investigating the ecological monitoring is investigating and live as adults along the coast. PISCO’s PISCO’s as adults along the coast. and live and of oceanographic long-term program ing into adults. Ocean currents strongly strongly currents Ocean ing into adults. and when barnacles settle where affect ing themselves with adhesive and matur with adhesive ing themselves plius stage to the cyprid stage. Eventually, Eventually, plius stage to the cyprid stage. attach surfaces, cyprids settle onto hard into the water. The young, or larvae, molt or larvae, The young, into the water. the nau from developing times, several ranging in size from 0.25 to 1 millimeter, 0.25 to 1 millimeter, ranging in size from adult and young life stages of barnacles. stages of barnacles. life adult and young young, free-swimming The adults release The illustration above shows generalized shows The illustration above species to gain insight into the ecology of larvae. with free-swimming invertebrates hard surfaces, where they live as adults. as adults. live they where surfaces, hard Ecologists often study barnacles as model months, they attach onto rocks and other attach onto rocks they months, are carried by ocean currents, possibly possibly ocean currents, by carried are or a period of weeks After long distances. invertebrates begin life as tiny larvae tiny as begin life that invertebrates Barnacle Life History other marine and many mussels, Barnacles, Year Position along coast Position

Kelp canopy biomass (tons/kilometer of coast)

PISCO scientists at UCSB and UCSC analyzed data from monthly aerial surveys to produce this figure showing changes in kelp beds in southern California over 34 years. Green indicates abundant kelp. Large kelp beds characterize some sections along the coast, such as at sites A, B, and C, but the amount of kelp at any site varied over time. At site C, for example, the strong El Niño in 1997-1998 caused a die-off of kelp. Survey data from ISP Alginates, Inc. ecological linkages Long-term Changes in Kelp Forests

nderwater forests of giant kelp are extremely important habitats along the U.S. West Coast, A large kelp forest in the Channel Islands is visible from the air. Photo: Ben Waltenberger U supporting numerous fish and invertebrate species. El Niño and storms strongly affect the Dispersal of Kelp Spores locations and sizes of kelp forests by removing kelp To reproduce, kelp release spores that from the sea bottom, which in turn influences the drift some distance away before settling animals that live there. onto the sea bottom. Knowing typical dispersal distances of kelp spores is critical To understand how climate change might affect kelp forests, PISCO scientists to understanding regional changes in kelp collaborated with UCSB researcher Dan Reed on a project funded by the forests. PISCO researchers in collabora- National Science Foundation to investigate the long-term trends in giant kelp tion with UCSB biologist Dan Reed have abundance along 500 kilometers of the southern California coast over 34 created a mathematical model that incor- years. Together, they combined monitoring data, oceanographic modeling, porates coastal oceanography and spore and field experiments to estimate extinction and colonization rates of kelp biology to mimic spore dispersal. Predic- patches in relation to oceanographic conditions, patch size, and connectivity tions from the model compared favorably with other kelp forests. to real-world measurements of spore Data from monthly aerial surveys showed that patches of giant kelp under- dispersal from an experimental kelp popu- went frequent extinctions and recolonizations over time scales from several lation. The model allows the scientists to months to over a decade (see figure, opposite page). In most cases, kelp examine the importance of factors such as forests disappeared for less than two years before being recolonized by new 11 wave height, current speed, turbulence, and kelp. Small, isolated kelp forests were more likely to go extinct than were large height at which the spores are released forests surrounded by other forests. Conversely, recolonization was more likely above the sea bottom in affecting spore for large patches and less likely for isolated patches. dispersal patterns. The research shows The scientists also studied the dispersal of kelp spores, which enable the kelp that while many kelp spores settle close to recolonize the sea bottom (see sidebar). This investigation of kelp forest to their parent, a sizeable fraction travel dynamics provides new insights into the degree of connectivity among kelp hundreds to several thousands of meters forests and tools for better predicting effects of changing ocean climate. before settling on the seafloor. While shorter-distance dispersal is adequate for the maintenance of extant kelp forests, longer-distance dispersal is needed for the recolonization of kelp in forests that have gone locally extinct.

Kelp researchers are Dan Reed (UCSB); Brian Gaylord, Brian Kinlan, and Libe Washburn (PISCO/UCSB); and Peter Raimondi and Patrick Drake (PISCO/UCSC). Brian Gaylord is now an assistant professor at the University of California, Davis.

Publications: Physical-biological coupling in spore dispersal of kelp forest macroalgae. Journal of Marine Systems 49 (2004): 19-39. Macroalgal spore dispersal in coastal environ- ments: Mechanistic insights revealed by theory and experiment. Ecological Monographs (in review). A metapopulation perspective on patch dynamics and connectivity of giant kelp. In J.P. Kritzer and P.F. Sale (eds.). Marine Metapopulations. 2006. Academic Press, San Diego. Kelp rockfish (Sebastes atrovirens) in a kelp forest (Macrocystis pyrifera). Photo: Luke Miller

0)3#/#OASTAL#ONNECTIONS„6OLUME From 1999 to 2005, PISCO measured many ecological parameters at multiple sites at each cape indicated on the map. Patterns found in four parameters are illustrated above. Abun- dance of macrophytes (seaweeds and seagrasses) correlated inversely with invertebrate abundance at many capes. At Cape Foulweather in Oregon, for example, macrophytes were abundant, and invertebrates were scarce. Concentrations of the important nutrient nitrate were lower in the intermittent upwelling regime than in the persistent upwelling regime, and 12 phytoplankton were more abundant. Cape Perpetua consistently had low nitrate concentrations and high chlorophyll concentrations.

Regional Oceanography Shapes Ecological Patterns Ecological Monitoring Ecological studies of marine systems traditionally focus on local biological inter- A major component of PISCO is a coor- actions such as predation and competition. However, large-scale events like El dinated, large-scale program of ecological Niño or climate change can alter local ecology by affecting supplies of larvae, and oceanographic monitoring along the nutrients, and phytoplankton. In multi-year studies along 1,300 kilometers of U.S. West Coast. The monitoring program the coast, PISCO scientists are finding that ecological patterns on the scale of includes: capes and bays can arise from interactions between regional-scale oceano- • measuring population replenishment of graphic events and local-scale ecological processes. key shoreline invertebrates and reef fish In northern California and Oregon, two distinct oceanographic regimes occur. from spring to late fall, North of Cape Blanco, Oregon, the strong jet of the California Current flows • biological surveys at approximately 50 parallel and close (20-50 kilometers) to shore, and summertime upwelling of rocky reef and kelp forest sites and 100 cold water is sporadic; this regime is called “intermittent upwelling.” South of shoreline sites, and Cape Blanco, the jet meanders as far as 300 kilometers offshore, and upwell- • monthly coastal transects in Oregon to ing is more consistent; this regime is called “persistent upwelling.” monitor conditions at representative sites. Spanning both regimes, PISCO scientists have measured numerous ecological Through these integrated monitoring parameters. The results show clear patterns, such as relatively high chlorophyll efforts, PISCO identifies changes in the concentrations and lower nutrient concentrations in the intermittent upwell- ecosystem and designs experiments to ing regime and inverse relationships between macrophyte and invertebrate understand the underlying causes. More abundance. Researchers are now evaluating the extent to which large-scale than six years of large-scale monitoring oceanographic conditions determine differences in the patterns and underly- data from PISCO provides scientists and ing processes in ecological communities, such as population replenishment resource managers with detailed informa- rates, organism growth, and species interactions. tion on species distributions, population PISCO/OSU researchers are Bruce Menge, Francis Chan, Sally Hacker, Maria fluctuations, shifts over time, and oceano- Kavanaugh, and Christopher Krenz. Maria Kavanaugh is now a doctoral stu- graphic changes along the coast. dent at the College of Oceanic and Atmospheric Sciences at OSU. Christopher For more information: www.piscoweb.org Krenz is now a John A. Knauss Marine Policy Fellow in Washington, D.C.

0ARTNERSHIPFOR)NTERDISCIPLINARY3TUDIESOF#OASTAL/CEANS ecological linkages ecological

Effects of Water Temperature on Urchin Survival Sea urchins develop from fertilized egg to larva in an uncertain world, drift- ing with the ocean currents. Prevailing oceanographic conditions can expose these vulnerable life stages to new environmental conditions and transport them away from their adult habitat. PISCO/UCSB scientists are studying the effects of temperature changes on the pluteus larvae of four common ur- chins in the genus Strongylocentrotus. The goal is to understand the role of water temperature in setting the northern and southern limits of the species’ ranges along the west coast. The scientists have found that differences as small as one degree can have dramatic effects down to the molecular level. Interestingly, urchin larvae from colder, northern waters are more sensitive to changes in temperature than their southern relatives. The geographic range of northern urchins likely is limited by temperature at its southern end. Know- ing how urchin larvae respond to changing temperatures at large geographic scales may enable predictions about the impacts of both short- and long-term climate changes on the distribution and abundance of these ecologically and commercially important species. For example, northern species are likely to be more susceptible to ocean warming than their southern relatives. PISCO/UCSB researchers are Gretchen Hofmann, LaTisha Hammond, and Kevin Fielman. Map (above): White and green sea urchins occur from southern Puget Sound to Alaska. Purple and red sea urchins are found from Baja California to Alaska.

Graph (right): Percent mortality of the young (pluteus larvae) of four urchin species exposed to different tem- peratures in the laboratory. Young red and purple urchins tolerated high temperatures better than white and green urchins, which inhabit colder, northern waters.

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Are Intertidal and Subtidal Species at Equal Risk? Climate warming is expected to have profound effects on the ocean. Research by PISCO/Stanford scientists shows that some of the ecological outcomes may seem paradoxical. The researchers tested the physiological tolerances of intertidal and subtidal snail species and determined the snails’ capacities for acclimating to warmer temperatures. A surprising finding was that intertidal snail species, despite tolerating higher temperatures than their subtidal rela- tives, face a greater threat from climate warming. Two factors account for this result. First, the intertidal species currently experience temperatures closer to their limits of heat tolerance. Second, they have more limited abilities to accli- mate to higher temperatures. Consequently, intertidal snails are vulnerable to The body temperature of the shoreline-dwelling snail the warming climate, while subtidal species have the capacity to tolerate some Tegula funebralis (green line) sometimes went above the temperature at which its heart function was impaired warming. (green horizontal line). The body temperature of the deeper-water snail T. brunnea (blue line) did not cross PISCO/Stanford researchers are Emily Stenseng, Caren Braby, and George the threshold (blue horizontal line). This result indicates Somero. Caren Braby is now a postdoctoral fellow at the Monterey Bay Aquar- that climate warming threatens T. funebralis more than T. brunnea. Body temperature fluctuation data from Tomanek ium Research Institute. and Somero (1999). Publication: Evolutionary and acclimation-induced variation in the thermal limits of heart function in congeneric marine snails: implications for vertical zonation. Biological Bulletin 208 (2005): 138-144.

0)3#/#OASTAL#ONNECTIONS„6OLUME Interdisciplinary Courses Train the Next Generation

n 2005, PISCO offered two interdisciplinary courses in marine research and policy as part of its training program for the next generation of scientists. The participants hailed from the four PISCO campuses— I UCSB, UCSC, Stanford, and OSU—and twelve other institutions. PISCO’s new Marine Conservation Science and Policy graduate course was held at Hatfield Marine Science Center in Newport, Oregon. PISCO scientists Jane Lubchenco and Steve Gaines taught this course with marine policy experts Andy Rosenberg (University of New Hampshire) and David Festa (Environmental Defense). PISCO policy coordinators Liz Riley, Satie Airamé, and Cinamon Vann co-instructed and provided logistical support. The intensive ten-day course engaged students in hands-on, interactive learning about the political and legal aspects of marine management, the role of science in marine policy, and the communication of science to wider audiences. “There is nothing else like this course in terms of quality and its unique combination of topics and approach,” said one participant. In 2005, twenty students attended PISCO’s month-long Biomechanics, Ecological Physiology, and Genetics of Intertidal Communities course at Stanford University’s Hopkins Marine Station. PISCO scientists Mark Denny, Steve Palumbi, and George Somero taught the course. “It was an absolutely fantastic experience,” said one participant. “I learned some good practical skills, [and] I’m thinking about ecology in very different ways.” interdisciplinary training& research interdisciplinary training & research interdisciplinary

Student Showcase

Does Detritus Determine Differences?

PISCO/UCSC doctoral student Jared Figurski is in- vestigating the ecological significance of kelp detritus, a rich mix of decaying kelp and microorganisms. He has found that reefs along the central coast of California vary greatly in the quantity of detritus. Because clumps of detritus are important as food and shelter for fish and other animals, Figurski is testing the hypothesis that the abundance of detritus leads to ecological differences among kelp forests. In 2004, his experiments showed that young-of-the-year rockfish actively seek out patches of kelp detritus as a nursery habitat. In 2005, he studied the importance of kelp detritus as habitat and food for fish, shrimp, PISCO/UCSC student Jared Figurski. Photo: Wyatt Patry amphipods, crabs, sea stars, and snails.

Physiology and Ecology of Mussel Reproduction

PISCO/OSU doctoral student Laura Petes is studying the effects of environmental stressors on the reproduction of mussels on rocky shores. She found that mussels at the upper edge of the mussel bed, an area of high heat stress, grow more slowly, allocate less energy towards reproduc- tion, and spawn earlier than do mussels in the lower edge of the mussel bed. She also has discovered new patterns of pigmentation in mussel reproductive tissue. Whereas males traditionally have been identified by white reproductive tissue and females by orange tissue, Petes found that both male and female 15 mussels in the upper edge of the mussel bed have orange reproductive tis- sue. The orange color arises from carotenoid pigments, which protect against damage by oxygen free radicals. Petes is testing whether the high carotenoid content helps to protect gametes from heat stress.

PISCO/OSU student Laura Petes. Photo: Jane Lubchenco

Using Genetics to Explore Ecological Linkages

PISCO/UCSB doctoral student Kim Selkoe used genet- ic techniques to examine possible interdependence of fish populations across the U.S.-Mexican border. She studied kelp bass (Paralabrax clathratus), which supports a large recreational fishery in southern California and an artisanal fishery in Baja California. Many scientists and policy-makers believe that south- ern California populations of kelp bass are replenished during strong El Niño years by young fish spawned in Baja and carried north by currents. However, Selkoe found that kelp bass of southern California have very different genetic traits and higher genetic diversity than those of Baja. Her findings indicate that kelp bass populations in California are self-sustaining, not dependent on Baja populations. Kim Selkoe is now a postdoctoral researcher at the Hawaii Institute of Marine Biology. Former PISCO/UCSB doctoral student Kim Selkoe (left) with fishing assistant Merit McCrea. Photo: Roland Takayama

Opposite page photos, left to right: Mike Nish, Jane Lubchenco, Satie Airamé

0)3#/#OASTAL#ONNECTIONS„6OLUME PISCO Teaches Monitoring Methods to Mexican Fishermen

ISCO partnered with the Mexican nonprofit Comunidad y Biodiversidad (Community and Biodiversity, or COBI) and a local fishing cooperative in August 2005 to P initiate monitoring of subtidal biological communities in no- take marine reserves near Isla Natividad on Mexico’s Pacific coast. Members of the cooperative decided to close portions of their fishing area for six years, with a plan to evaluate the effects and possibly increase the size of the marine reserve system later. Scientists and the fishermen worked together to identify local species of commercial and ecological importance. These species will be monitored regularly, both in areas closed to fishing and those that are fished, to determine the effects of the no-take reserves. This collaboration represents the first of many under a newly formed project focusing on small-scale fishing cooperatives in Mexico. PISCO/UCSC technician Amanda Jensen traveled to Mexico to work with COBI and the fishermen to modify PISCO’s protocols to enable the local fishermen, often using hookah diving apparatus, to monitor changes in key species. For more information about PISCO’s subtidal monitoring program, including diver training, visit www.piscoweb.org.

sharing THE science sharing the science

PISCO Advises California on Marine Protection Science

The California Resources Agency is working to implement the Marine Life Protection Act (MLPA) enacted by California in 1999. The agency selected the central coast from Point Conception to Pigeon Point for the first phase of a network of marine protected areas (MPAs). PISCO scientists Steve Gaines, Mark Carr, and Steve Palumbi serve on a Science Advisory Team, sharing their scientific knowledge through presentations to MLPA decision- makers and stakeholders on topics from larval dispersal to MPA network design. The scientists helped to identify species that may benefit from MPAs, and Carr met regularly with stakeholders from the central coast region. PISCO policy coordinators Satie Airamé and Cinamon Vann interact closely with state and federal officials, fishermen, and environmental groups to ensure that PISCO’s research findings and data are available to answer questions and inform policy. For more information on the MLPA, go to www.dfg.ca.gov/mrd/mlpa.

Partington Point, near Big Sur, is one area that will be considered for protection under the MLPA. Photo: Haven Livingston

Comparing Methods of Stock Assessment In 2003 and 2005, PISCO worked with commercial live-fish fishermen and the California Department of Fish and Game 17 to compare two methods of assessing fish stocks. Accurate information about the stocks is crucial for setting sustainable catch limits and evaluating marine pro- PISCO scientists Steve Gaines and Steve Palumbi work with Monterey Bay National Marine Sanctuary representative tected areas. In Carmel Bay, the scientists Holly Price during MLPA proceedings. Photo: Satie Airamé and fishermen compared estimates of fish abundance from visual surveys by Straight Talks on Fishing scuba divers with those estimated from catch-per-unit-effort (CPUE) by fishing. As Following on the success of the first series of Straight Talks, expected, divers counted more fish species a second series is being organized by PISCO, California Sea Grant, and (38) than were caught by fishing gear (20), representatives of the central California fishing community. The Straight Talk and fish abundance and number of species forums bring together scientists and fishermen to share their knowledge of varied with fishing gear type and method the coastal ocean environment and fishing. PISCO/UCSC scientist Mark Carr of deployment. One key result was that and Sea Grant Advisor Rick Starr will lead discussions as they did in the initial some fish species were sampled better by Straight Talks. The forums create an informal, congenial setting where people fishing (e.g., cabezon, grass rockfish), while can freely discuss topics that often are contentious. other species were sampled better by div- Coastal Management in Oregon ers (e.g., kelp rockfish, kelp greenling). Find- ings from the 2003 and 2005 studies will PISCO is participating in an initiative to be used to improve fish survey programs update Oregon’s Rocky Shore Management by integrating fishing and diver surveys for Strategy, which was adopted in 1994 and more comprehensive assessments. provides a framework for coastal manage- ment, focusing on 39 rocky shore sites. The PISCO/OSU policy coordinator sits on the Technical Advisory Committee that guides the project, PISCO staff compiled data sets for decision-makers, and PISCO field

Opposite page photos: Matt Robart (left), researchers are monitoring human use of Luis Bourillón (right) Cape Meares, Oregon. Photo: Matt Robart Oregon’s rocky shores.

0)3#/#OASTAL#ONNECTIONS„6OLUME Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO)

For more information: Web site: www.piscoweb.org E-mail: [email protected]

PISCO Oregon State University Department of Zoology 3029 Cordley Hall Corvallis, OR 97331

PISCO University of California, Santa Cruz Long Marine Laboratory 100 Shaffer Road Santa Cruz, CA 95060

PISCO University of California, Santa Barbara Marine Science Institute Santa Barbara, CA 93106-6150

PISCO Stanford University Hopkins Marine Station Oceanview Boulevard Pacific Grove, CA 93950

Photos, top to bottom and left to right: Roly Russell, Gretchen Hofmann, Monica Pessino, Jane Lubchenco, Luke Miller, Sean Hoobler, Monica Pessino, Jared Figurski, Gil Rilov, Jared Figurski, Cristine McConnell, Gil Rilov, Tui Anderson, Luke Miller, Cristine McConnell, Haven Livingston

Paper stock contains 50% recycled content, 15% post-consumer content. Printed with linseed oil-based inks.