Kristy KroekerPost Doctoral Researcher
My research focuses on investigating the impacts of environmental change on coastal marine ecosystems. More information about my research >

University of California, Davis – Department of Evolution and Ecology
Professor, Bodega Marine Laboratory
Bodega Marine Laboratory
P.O. Box 247
Bodega Bay , CA 94923
Climate Change, Biogeography, and Local Adaptation
The Sanford Lab is interested in how marine populations and communities vary in response to both natural oceanographic variation and anthropogenic climate change. Our research seeks to synthesize the ecological, evolutionary, physiological, and oceanographic processes that shape marine populations: both over large distances along coastlines, and in an era of accelerating climate change. We seek mechanistic understanding of these processes through coordinated field and laboratory experiments centered at Bodega Marine Laboratory. Much of our work focuses on marine intertidal communities, where organisms and their interactions are diverse and easily studied.
Ongoing research projects lie primarily in three areas:
For additional information see Research tab, above.

The Sanford Lab (Summer 2011): Back row: Kirk Sato (technician), Cristian Monaco (visiting graduate student), Dan Swezey (graduate student), Evelyne Kuo (graduate student), Eric Sanford; Front row: Morgan Kelly (graduate student), Beth Lenz (technician), Megan Young (technician). Not shown: Jill Bible (graduate student).
Marine species are often distributed over thousands of kilometers of coastline and thus separate populations can experience strikingly different environments. However, we know surprisingly little about the extent to which environmental variation shapes evolutionary differences among populations of marine species. Our recent experiments have documented regional variation in the capacity of a predatory snail (Nucella canaliculata) to drill thick-shelled mussels. These differences in drilling capacity have a genetic basis and appear to reflect local adaption to variation in prey recruitment in California versus Oregon. Our results suggest that geographic mosaics of selection imposed by persistent oceanographic variation can shape adaptive differentiation among populations of marine species in adjacent coastal regions. We are continuing to combine field studies, laboratory experiments, and analyses of molecular markers (microsatellites) to investigate (1) the scale at which adaptive differentiation occurs in marine species, and (2) the ecological consequences of these patterns, in a variety of organisms including snails, tidepool copepods, bryozoans, and oysters.

Dogwhelk, mussels, and egg capsules (left), and dogwhelks feeding on prey (right).
Selected publications:
Kelly, M.W., E. Sanford, and R.K. Grosberg. 2011. Limited potential for adaptation to climate change in a broadly-distributed marine crustacean. Proceedings of the Royal Society of London: Biological Sciences Series B, doi: 10.1098/rspb.2011.0542.
Sanford, E. and M.W. Kelly. 2011. Local adaptation in marine invertebrates. Annual Review of Marine Science 3: 509–535.
Sanford, E. and D.J. Worth. 2010. Local adaptation along a continuous coastline: prey recruitment drives differentiation in a predatory snail. Ecology 91: 891–901.
Sanford, E. and D.J. Worth. 2009. Genetic differences among populations of a marine snail drive geographic variation in predation. Ecology 90: 3108–3118.
Kuo, E.S.L. and E. Sanford. 2009. Geographic variation in the upper thermal limits of an intertidal snail: implications for climate envelope models. Marine Ecology Progress Series 388: 137–146.
Sanford, E., M.S. Roth, G.C. Johns, J.P. Wares, and G.N. Somero. 2003. Local selection and latitudinal variation in a marine predator-prey interaction. Science 300: 1135–1137.
Funding for this work is provided by the National Science Foundation under Grant No. OCE-06-22924.
What climatic and biological factors set geographic range limits and to what extent can these boundaries be overcome through adaptation and/or phenotypic plasticity? What ecological and evolutionary processes facilitate or impede range extensions? In New England, we have used larval rearing experiments and field studies to identify the factors that maintain the northern range limit of the mud fiddler crab (Uca pugnax) near Cape Cod, Massachusetts. In California, we are addressing related questions with the volcano barnacle (Tetraclita rubescens), a species that has undergone a range extension along the coast of northern California during the past 25 years. Other projects in the lab are investigating geographic range boundaries using the tidepool copepod Tigriopus californicus, and the limpet Lottia insessa (a specialist herbivore on the feather boa kelp Egregia menziesii).

Fiddler crab releasing larvae (left), and volcano barnacles (right).
Selected publications:
Dawson, M.N., R.K. Grosberg, Y.E. Stuart, and E. Sanford. 2010. Population genetic analysis of a recent range expansion: mechanisms regulating the poleward range limit in the volcano barnacle Tetraclita rubescens. Molecular Ecology 19: 1585–1605.
Sanford, E. and D.S. Swezey. 2008. Response of predatory snails to a novel prey following the geographic range expansion of an intertidal barnacle. Journal of Experimental Marine Biology and Ecology 354: 220–230.
Sanford, E., S.B. Holzman, R.A. Haney, D.M. Rand, and M.D. Bertness. 2006. Larval tolerance, gene flow, and the northern geographic range limit of fiddler crabs. Ecology 87: 2882–2894.
Thermal tolerance, species interactions, and climate change
Many predictions have focused on how climate change might impact species through the direct effects of environmental stress on demographic rates. These models treat species as independent units and often neglect the fact that organisms are embedded within complex webs of interacting species. We are intrigued by the possibility that some of the most immediate and important impacts of climate change could arise through changes in key species interactions. Using field and laboratory experiments, we have shown that the effect of a keystone predator, the sea star Pisaster ochraceus, is influenced by small changes in ocean temperature (~3°C). Thus, long-term shifts in cold-water upwelling patterns could generate community-level effects through impacts on this keystone predator. More recently, we have also investigated how the feeding and growth of Pisaster is influenced by exposure to aerial conditions during low tide. In other projects, we are examining the extent to which populations of marine species (including intertidal snails and copepods) are locally adapted to biogeographic variation in temperature. This work includes the use of selection experiments to test the capacity of tidepool copepods to adapt to future increases in temperature.

Sea stars (Pisaster ochraceus) feeding on mussels.
In collaboration with Drs. Brian Gaylord, Tessa Hill, and Ann Russell, we are examining the influence of ocean acidification on ecologically and economically important species in northern California (with a focus on the Sonoma coast and Tomales Bay). Our interdisciplinary research combines moored and shipboard measurements of seawater chemistry with laboratory and field studies of the biological effects of ocean acidification. To date, our experiments on the native oyster (Ostrea lurida) and California mussel (Mytilus californianus) indicate that larvae and juveniles of these important foundation species may be quite vulnerable to decreasing pH and changes in the calcium carbonate saturation state. See Bodega Ocean Acidification Research (BOAR)
Bodega Ocean Acidification Research (BOAR) in the News:
Sept 14, 2011 KQED-TV, QUEST: Will ocean acidification harm sea urchins?
July 18, 2011 KQED Radio: Climate Change Threatens California Mussels
July 18, 2011 The Orange County Register: California mussels: 1st warming casualty?
July 15, 2011 KQED News ClimateWatch: Study: Climate Change Muscling in on Mussels
July 14, 2011 UC Davis News and Information: Acid oceans could hit California mussels
July 6, 2010 KGO-TV/ABC News: Oysters could hold key to ocean acidification.
May 22, 2010 KNTV/NBC News: BML researchers study the effects of ocean acidification on Tomales Bay oysters
April 22, 2010 MSNBC.com: Acidic oceans worsening, experts warn - CO2 impact coming faster than seas can adapt, they say
April 19, 2010 National Science Foundation News: On 'Earth Week', World Is No Longer Our Oyster - Acidifying oceans dramatically stunt growth of already threatened shellfish

System at BML to manipulate CO2 in larval cultures (left), and magnified view of oyster larvae (right).
Our BOAR team is also part of a broader consortium called the Ocean Margin Ecosystems Group for Acidification Studies (OMEGAS). OMEGAS is an interdisciplinary consortium of researchers from 7 institutions studying ocean acidification along the coasts of California and Oregon.
The location of Bodega Marine Laboratory within a major upwelling center, and the lab's research strengths in coastal oceanography (Bodega Ocean Observing Node), make this an ideal place to explore links between oceanographic processes and the dynamics of benthic marine communities. In addition to understanding the effects of changing temperature and ocean chemistry, our lab continues to be interested in how variation in bottom-up forces affects invertebrate reproduction, recruitment, and growth.
Selected publications:
Gaylord B., T.M. Hill, E. Sanford, E.A. Lenz, L.A. Jacobs, K.N. Sato, A.D. Russell, and A. Hettinger. 2011. Functional impacts of ocean acidification in an ecologically critical foundation species. Journal of Experimental Biology 214: 2586–2594.
Kelly, M.W., E. Sanford, and R.K. Grosberg. 2011. Limited potential for adaptation to climate change in a broadly-distributed marine crustacean. Proceedings of the Royal Society of London: Biological Sciences Series B, doi: 10.1098/rspb.2011.0542.
Pincebourde, S., E. Sanford, and B. Helmuth. 2009. An intertidal sea star adjusts thermal inertia to avoid extreme body temperatures. American Naturalist 174: 890–897.
Kuo, E.S.L. and E. Sanford. 2009. Geographic variation in the upper thermal limits of an intertidal snail: implications for climate envelope models. Marine Ecology Progress Series 388: 137–146.
Pincebourde, S., E. Sanford, and B. Helmuth. 2008. Body temperature during low tide alters the feeding performance of a top intertidal predator. Limnology and Oceanography 53: 1562–1573.
Sanford, E. 2002. Water temperature, predation, and the neglected role of physiological rate effects in rocky intertidal communities. Integrative and Comparative Biology 42: 881–891.
Sanford, E. and B.A. Menge. 2001. Spatial and temporal variation in barnacle growth in a coastal upwelling system. Marine Ecology Progress Series 209: 143–157.
Sanford, E. 1999. Regulation of keystone predation by small changes in ocean temperature. Science 283: 2095–2097.
Teaching on the UC Davis main campus:
EVE 112/112L. Biology of Invertebrates (Winter 2010 and alternate years; co-taught with Rick Grosberg.) Survey of the major invertebrate phyla focusing on form and function, ecology, and phylogenetic relationships. Are you intrigued by corals, octopus, barnacles, and sea urchins? This is the course for you! (3-unit lecture course; 2-unit lab that emphasizes study of live animals, 2 field trips)
EVE 101. Introduction to Ecology (co-taught in alternate years.) A survey of the general principles of ecology. (4-unit lecture/discussion course)

Invertebrates from the California coast: Giant green sea anemone (left), and sunflower sea star (right).
Teaching at the Bodega Marine Laboratory:
Note: These courses are offered at the coast each summer. Please consult the BML student information web pages for details on housing and applications.
EVE 114. Experimental Invertebrate Biology.Want to learn more about the remarkable diversity of tidepool animals that make their home on the rugged northern California coast? This is the course that you have been looking for! We will cover the biology, ecology, and evolution of local marine invertebrates with a focus on adaptations to environmental and biological factors encountered on the California coast. This course offers hands-on field and laboratory components with an emphasis on testing hypotheses that we generate as a class. Short class projects provide students with practical experience in all aspects of the scientific process including making observations, generating hypotheses, designing experiments, collecting and analyzing data, and scientific writing (3-units lecture/field/lab).
This course is offered each year during Summer Session I. Note that students often take this course at the same time as Brian Gaylord’s Mechanical Design in Organisms (EVE 106). These two courses complement each other very well.
BIS 124. Coastal Marine Research. In this 3-unit course, students pursue independent research projects related to either EVE 106 (Mechanical Design in Organisms) or EVE 114 (Experimental Invertebrate Biology). You will receive training in all phases of the scientific process from experimental design to data analysis to communication of results.
EVE 111. Marine Environmental Issues (co-taught with Brian Gaylord) This 1-unit course is built around readings and informal discussions related to marine conservation and major environmental issues in coastal waters. Topics include the impacts of climate change, invasive species, and overfishing.
Please email Eric Sanford for pdf reprints.
Gaylord B., T.M. Hill, E. Sanford, E.A. Lenz, L.A. Jacobs, K.N. Sato, A.D. Russell, and A. Hettinger. 2011. Functional impacts of ocean acidification in an ecologically critical foundation species. Journal of Experimental Biology 214: 2586–2594.
Kelly, M.W., E. Sanford, and R.K. Grosberg. 2011. Limited potential for adaptation to climate change in a broadly-distributed marine crustacean. Proceedings of the Royal Society of London: Biological Sciences Series B, doi: 10.1098/rspb.2011.0542.
Sanford, E. and M.W. Kelly. 2011. Local adaptation in marine invertebrates. Annual Review of Marine Science 3: 509–535.
Kelly, M.W., and E. Sanford. 2010. The evolution of mating systems in barnacles. Journal of Experimental Marine Biology and Ecology 392: 37–45.
Dawson, M.N., R.K. Grosberg, Y.E. Stuart, and E. Sanford. 2010. Population genetic analysis of a recent range expansion: mechanisms regulating the poleward range limit in the volcano barnacle Tetraclita rubescens. Molecular Ecology 19: 1585-1605.
Sanford, E. and D.J. Worth. 2010. Local adaptation along a continuous coastline: prey recruitment drives differentiation in a predatory snail. Ecology 91: 891-901.
Sanford, E., M.E. Wood, and K.J. Nielsen. 2009. A non-lethal method for estimation of gonad and pyloric caecum indices in sea stars. Invertebrate Biology 128: 372-380.
Pincebourde, S., E. Sanford, and B. Helmuth. 2009. An intertidal sea star adjusts thermal inertia to avoid extreme body temperatures. American Naturalist 174: 890-897.
Sanford, E. and D.J. Worth. 2009. Genetic differences among populations of a marine snail drive geographic variation in predation. Ecology 90: 3108-3118.
Kuo, E.S.L. and E. Sanford. 2009. Geographic variation in the upper thermal limits of an intertidal snail: implications for climate envelope models. Marine Ecology Progress Series 388: 137-146.
Sanford, E. and M.D. Bertness. 2009. Latitudinal gradients in species interactions. Ch. 14 in: J.D. Witman, J.D. and K. Roy (eds), Marine Macroecology. University of Chicago Press, Chicago. In press.
Pincebourde, S., E. Sanford, and B. Helmuth. 2008. Body temperature during low tide alters the feeding performance of a top intertidal predator. Limnology and Oceanography 53: 1562-1573.
Sanford, E. and D.S. Swezey. 2008. Response of predatory snails to a novel prey following the geographic range expansion of an intertidal barnacle. Journal of Experimental Marine Biology and Ecology 354: 220-230.
Sanford, E. and B.A. Menge. 2007. Reproductive output and consistency of source populations in the sea star Pisaster ochraceus. Marine Ecology Progress Series 349: 1-12.
Menge, B.A., B.A. Daley, E. Sanford, E. P. Dahlhoff, and J. Lubchenco. 2007. Mussel zonation in New Zealand: Towards an integrative eco-physiological approach. Marine Ecology Progress Series 345: 129-140.
Sanford, E. 2007. Sea stars. Pp 505-509 in: Denny, M.W. and S.D. Gaines (eds), Encyclopedia of Tidepools and Rocky Shores, University of California Press, Berkeley.
Sanford, E., S.B. Holzman, R.A. Haney, D.M. Rand, and M.D. Bertness. 2006. Larval tolerance, gene flow, and the northern geographic range limit of fiddler crabs. Ecology 87: 2882-2894.
Sanford, E., M. S. Roth, G. C. Johns, J. P. Wares, and G. N. Somero. 2003. Local selection and latitudinal variation in a marine predator-prey interaction. Science 300: 1135-1137.
Tomanek, L. and E. Sanford . 2003. Heat-shock protein 70 (Hsp 70) as a biochemical stress indicator: An experimental field test in two congeneric intertidal gastropods (Genus: Tegula ). Biological Bulletin 205: 276-284.
Sanford, E. 2002. Water temperature, predation, and the neglected role of physiological rate effects in rocky intertidal communities. Integrative and Comparative Biology 42: 881-891.
Sanford, E. 2002. The feeding, growth and energetics of two rocky intertidal predators ( Pisaster ochraceus and Nucella canaliculata ) under water temperatures simulating episodic upwelling. Journal of Experimental Marine Biology and Ecology 273(2): 199-218.
Sanford, E. 2002. Community responses to climate change: links between temperature and keystone predation in a rocky intertidal system. In S.H. Schneider and T.L. Root (eds.), Wildlife Responses to Climate Change: North American Case Studies , pp.165-200. Island Press, Covelo, CA.
Menge, B.A., E. Sanford , B.A. Daley, T.L. Freidenburg, G. Hudson, and J. Lubchenco. 2002. An inter-hemispheric comparison of bottom-up effects on community structure: insights revealed using the comparative-experimental approach. Ecological Research 17(1): 1-16.
Sanford, E. and B.A. Menge. 2001. Spatial and temporal variation in barnacle growth in a coastal upwelling system. Marine Ecology Progress Series 209: 143-157.
Sanford, E. 1999. Regulation of keystone predation by small changes in ocean temperature. Science 283: 2095-2097.
Menge, B.A., B.A. Daley, J. Lubchenco, E. Sanford , E. Dahlhoff, P.M. Halpin, G. Hudson, and J.L. Burnaford. 1999. Top-down and bottom-up regulation of New Zealand rocky intertidal communities. Ecological Monographs 69(3): 297-330.
Miner, B.G., E. Sanford , R.R. Strathmann, B. Pernet, and R.E. Emlet. 1999. Functional and evolutionary implications of opposed bands, big mouths, and extensive oral ciliation in larval Opheliids and Echiurids (Annelida). Biological Bulletin 197(1): 14-25.
Menge, B.A., B. Daley, P.A. Wheeler, E. Dahlhoff, E. Sanford , and P.T. Strub. 1997. Benthic-pelagic links in rocky intertidal communities: evidence for bottom-up effects on top-down control. Proceedings National Academy of Sciences 94: 14530-14535.
Sanford, E., D. Bermudez, M.D. Bertness, and S.D. Gaines. 1994. Flow, food supply and acorn barnacle population dynamics. Marine Ecology Progress Series 104: 49-62.
Bertness, M.D., S.D. Gaines, D. Bermudez, and E. Sanford . 1991. Extreme spatial variation in the growth and reproductive output of the acorn barnacle Semibalanus balanoides . Marine Ecology Progress Series 75: 91-100.
Eric Sanford
B.A., Biology, Brown University (1990)
Ph.D., Zoology, Oregon State University (1999)
Post-doctoral Fellow, Stanford University (1999–2002)
Research Associate, Brown University (2002–2004)
Assistant Professor, UC Davis (2005–2010)
Associate Professor, UC Davis (2010–present)
Kristy KroekerMy research focuses on investigating the impacts of environmental change on coastal marine ecosystems. More information about my research >
Daniel S. SwezeyI am broadly interested in global change and the ecology of the oceans in a high CO2 world. My dissertation research is focused on local adaptation and spatially variable organismal response to future ocean acidification. Numerous recent experiments have documented the potential impacts of increasing ocean acidity on calcifying organisms. However, these organismal responses often vary among different taxonomic groups and even among individuals in a single species. The evolutionary basis for this variability remains largely unexplored. Using bryozoans, a unique group of calcifying colonial invertebrates, I am investigating how coastal upwelling and patterns of gene flow may shape the response of populations to elevated pCO2 in the future. As a member of the REACH (Responding to Rapid Environmental Change) interdisciplinary IGERT training program at UC Davis, I am also interested in connecting ecological findings to practical conservation decision-making, and I have taken a leadership role in collaborative research projects in agricultural landscapes, invaded riparian habitat, coral reef systems, and the deep sea.
Jill BibleMy broad interests are in human impacts on ecosystems, ecosystem structure and functioning, conservation, and restoration ecology. I am particularly interested in the effects of climate change on nearshore marine ecosystems such as estuaries, rocky intertidal zones, kelp forests, and coral reefs. My current research focuses on whether populations of marine invertebrates are locally adapted to their environments and whether local adaptation will make them more or less vulnerable to anthropogenic stressors. Specifically, I am investigating whether populations of Olympia oysters, the only native oyster species on the West Coast, are locally adapted to their home estuaries. My work focuses on whether populations of oysters from three different California estuaries have evolved different responses to abiotic factors expected to change with climate change, such as temperature, salinity, and carbonate chemistry. In addition to conducting research, I also have a passion for teaching and communicating science to diverse audiences.
Evelyne Sui Ling Kuo (Ph.D. awarded 2012), Graduate Group in Ecology
Morgan Kelly (Ph.D. Awarded 2011)
David Worth (Research technician, 2007-2008)
Kirk Sato (Research technician, 2008-2010)
Beth Lenz (Research technician 2009-2011)
Megan Young (Research technician, 2010-2011)Also see:
Interested in joining the Sanford Lab?
I am looking for bright and enthusiastic students who are fascinated by the ecology and evolution of marine organisms. The Sanford Lab offers opportunities for students at all levels.
Undergraduate Students:
There are often research opportunities available for motivated UC Davis undergraduates interested in working in the Sanford Lab. The students who pursue these opportunities often do so after having taken one or more of my courses. If you have a solid academic record, enjoy working hard, and think you might like to get involved, feel free to contact me.
Graduate Students:
I welcome inquiries from prospective graduate students. The Sanford Lab is based full-time at Bodega Marine Laboratory (BML) and accepts students through either the Graduate Group in Ecology or the Population Biology Graduate Group. Entering students generally spend their first year on campus completing coursework and then move to the coast to become full-time residents at BML.
I enjoy working with students who share interests with me, but I am also committed to training students who are independent thinkers and creative scientists. Thus, I expect my students to develop and pursue an exciting thesis of their own design (with my input and encouragement, of course!).
Experimental field studies are the backbone of my research and I urge students to test hypotheses in the field whenever possible. This is easily done given our location within the Bodega Marine Reserve and our proximity to many other superb field sites along the California coast. I am also convinced of the power of complementary lab studies, and the outstanding seawater facilities at BML create opportunities for a variety of larval rearing and mesocosm experiments. I encourage students to take an integrative approach to their research and to seek training in other disciplines where appropriate. For example, I am very interested in how physiology and population genetics can inform ecological and evolutionary questions and I welcome students with interests in developing skills in these areas.
Sanford Lab (Summer 2008)

The Sanford Lab (Summer 2008): Pictured from left to right are Morgan Kelly (graduate student), Jennifer Hoey (REU student), Chris Kwan (UCD undergraduate), Eric Sanford, David Worth (research technician), Evelyne Kuo (graduate student).
Climate Change, Biogeography, and Local Adaptation
The Sanford Lab is interested in how marine populations and communities vary in response to both natural oceanographic variation and anthropogenic climate change. Our research seeks to synthesize the ecological, evolutionary, physiological, and oceanographic processes that shape marine populations: both over large distances along coastlines, and in an era of accelerating climate change. We seek mechanistic understanding of these processes through coordinated field and laboratory experiments centered at Bodega Marine Laboratory. Much of our work focuses on marine intertidal communities, where organisms and their interactions are diverse and easily studied.
Ongoing research projects lie primarily in three areas:
For additional information see Research tab, above.

The Sanford Lab (Summer 2011): Back row: Kirk Sato (technician), Cristian Monaco (visiting graduate student), Dan Swezey (graduate student), Evelyne Kuo (graduate student), Eric Sanford; Front row: Morgan Kelly (graduate student), Beth Lenz (technician), Megan Young (technician). Not shown: Jill Bible (graduate student).