Eric Sanford, Ph.D.

Eric Sanford

Unit
College of Biological Sciences
Evolution and Ecology
Bodega Marine Laboratory

Bodega Marine Laboratory
University of California Davis, Bodega Marine Laboratory, PO Box 247, 2099 Westshore Rd, Bodega Bay CA 94923
Bio

Research | Teaching | Lab | Prospective Students | Publications

Research Interests

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 integrate ecology, evolution, and biogeography to understand the processes that shape marine communities: 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.

research interests

 

Ongoing research projects lie primarily in three areas:

  • Local adaptation in marine species
  • The regulation of species’ geographic range limits
  • The influence of climate change and ocean acidification on marine communities

Local adaptation in marine species

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)
Above: 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.

The regulation of species' geographic range limits

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).
 

range limits
Above: Fiddler crab releasing larvae (left), and volcano barnacles (right).

Selected publications:

Sanford, E.  2013.  The Biogeography of Marine Communities. Ch. 7 in: M.D. Bertness, J. Bruno, B.R. Silliman, and J.J. Stachowicz (eds), Marine Community Ecology and Conservation. Sinauer Press, Sunderland, MA. In press.

Kuo, E.S.L., and E. Sanford. 2013. Distribution of the seaweed limpet Lottia insessa along the Pacific coast.  Pacific Science 67: 303–313.

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.

The influence of climate change and ocean acidification on marine communities

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 are 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.

seastars
Sea stars (Pisaster ochraceus) feeding on mussels.


Bodega Ocean Acidification Research (BOAR)

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. Our interdisciplinary research program combines moored and shipboard measurements of seawater chemistry with laboratory and field studies of the biological effects of ocean acidification. 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.

For more information, including recent press coverage, see Bodega Ocean Acidification Research (BOAR)

ocean acidification lab
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. Recent results from a selection experiment conducted at BML suggests that purple sea urchin populations possess standing genetic variation that may confer some resilience to ocean acidification. 

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:

Sanford, E., B. Gaylord, A. Hettinger, E.A. Lenz, K. Meyer, and T.M. Hill. 2014. Ocean acidification increases the vulnerability of native oysters to predation by invasive snails. Proceedings of the Royal Society of London: Biological Sciences Series B 281: 20132681

Pespeni, M.H., E. Sanford, B. Gaylord,  T. M. Hill,  J. D. Hosfelt,  H. Jaris, M. LaVigne, E. A. Lenz,  A. D. Russell, M. K. Young,  S. R. Palumbi.  2013.  Evolutionary change during experimental ocean acidification.  Proceedings National Academy of Sciences, doi: 10.1073/pnas.1220673110.

Hettinger, A., E. Sanford, T.M. Hill, A.D. Russell, K.N. Sato, J. Hoey, M. Forsch, H.N. Page, and B. Gaylord.  2012.  Persistent carry-over effects of planktonic exposure to ocean acidification in the Olympia oyster. Ecology 93: 2758–2768.

Kelly, M.W., E. Sanford, and R.K. Grosberg.  2012. Limited potential for adaptation to climate change in a broadly distributed marine crustacean. Proceedings of the Royal Society of London: Biological Sciences Series B 279: 349–356.  

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.

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

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 the 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)

Giant green sea anemone (left), and sunflower sea star (right)
Invertebrates from the California coast: Giant green sea anemone (left), and sunflower sea star (right).

 

Teaching at Bodega Marine Laboratory:

Note: These courses are offered at the coast each year during Summer Session I. Please consult Undergraduate Courses at BML for the latest details on courses, 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).

BIS 124. Coastal Marine Research. In this 6-unit course, students pursue independent research projects related to EVE 114 (Experimental Invertebrate Biology).  You will receive training and practice in all phases of the scientific process from experimental design to data analysis and science communication (including preparing a final written report and creative filmmaking for a general audience).  Must be taken concurrently with EVE 114.

EVE 111. Marine Environmental Issues. 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.

The Sanford Lab

sanford lab 2015
The Sanford Lab (Summer 2015): From left to right: Olivia Turnross (graduate student), Dan Swezey (graduate student), Jill Bible (graduate student), Chris Kwan (graduate student), Eric Sanford (PI), Jason Toy (technician), Emily Rivest (BOAR post-doc).

 

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)

Former Lab Members

  • Daniel S. Swezey Ph.D.
  • Ryan Jenkinson Ph.D., Joint Doctoral Program in Ecology
  • Jill Bible Ph.D., Graduate Group in Ecology
  • Olivia R. Turnross, Ph.D. student, Graduate Group in Ecology
  • Chris Kwan Ph.D., Joint Doctoral Program in Ecology
  • Emily Rivest Ph.D., Post Doc
  • Evelyne Sui Ling Kuo (Ph.D. awarded 2012), Graduate Group in Ecology. Current position: Earthwatch Institute, Asia Pacific Region Program Coordinator
  • Morgan Kelly (Ph.D. Awarded 2011). Current position: Assistant Professor, Louisiana State University
  • David Worth (Research technician, 2007-2008)
  • Kirk Sato (Research technician, 2008-2010). Current position: Graduate Student, Scripps Institution of Oceanography, UC San Diego
  • Beth Lenz (Research technician 2009-2011). Current position: Graduate Student, University of Hawaii, Hawaii Institute of Marine Biology
  • Megan Young (Research technician, 2010-2011). Current position: Biology and Chemistry Teacher
  • Kristy Kroeker (BOAR Postdoctoral Researcher, 2012-2014). Current Position: Assistant Professor UC Santa Cruz
  • Seth Miller (BOAR Postdoctoral Researcher, 2012-2013). Current position: Postdoctoral Fellow, Smithsonian Environmental Research Center, Maryland
  • Kelly Laughlin (Research Technician, 2012-2014)
  • Kaylee Griffith (Research Technician, 2014). Current position: Graduate Student, San Diego State University
  • Jason Toy (Research Technician, 2015). Current position: Teaching Assistant, Bermuda Institute of Ocean Sciences

Prospective Students

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.

Publications

Please email Eric Sanford for pdf reprints.

Maynard, A., J.M. Bible, M.H. Pespeni, E. Sanford, and T.G. Evans. 2018. Transcriptomic responses to extreme low salinity among locally adapted populations of Olympia oyster (Ostrea lurida).  Molecular Ecology, in press.

Goddard, J.H.R., N. Treneman, T. Prestholdt, C. Hoover, B. Green, W.E. Pence, D.E. Mason, P. Dobry, J.L. Sones, E. Sanford, R. Agarwal, G.R. McDonald, R.F. Johnson, and T.M. Gosliner. 2018. Heterobranch sea slug range shifts in the northeast Pacific Ocean associated with the 2015-16 El Niño. Proceedings of the California Academy of Sciences Series 4, Volume 65(3): 107–131.

Agha, M., M.K. Riley, E. Sanford, J.T. Carlton, W.A. Newman, and B.D. Todd. 2018.  A review of epizoic barnacles reported from freshwater turtles, with a new record from California. Herpetological Review 49(1): 25–28.

Swezey, D.S., J.R. Bean, T.M. Hill, B. Gaylord, A.T. Ninokawa, and E. Sanford. 2017.  Plastic responses of bryozoans to ocean acidification. Journal of Experimental Biology 220: 4399–4409.

Bible, J.M., B.S. Cheng, A.L. Chang, M.C. Ferner, K. Wasson, C.J. Zabin, M. Latta, E. Sanford, A. Deck, and E.D. Grosholz. 2017. Timing of climate-driven stressors alters interactive effects on an estuarine foundation species. Ecology 98(9): 2468–2478.  

Davis, C.V., E.B. Rivest, T.M. Hill, B. Gaylord, A.D. Russell, E. Sanford. 2017. Ocean acidification compromises the performance of planktonic foraminifera with implications for carbon cycling. Scientific Reports, doi: 10.1038/s41598-017-01530-9

Chan, F., J. A. Barth, C. A. Blanchette, R. H. Byrne, F. Chavez, O. Cheriton, R.A. Feely, G. Friederich, B. Gaylord, T. Gouhier, S.Hacker, T. Hill, G. Hofmann, M.A. McManus, B.A. Menge, K.J. Nielsen, A. Russell, E. Sanford, J. Sevadjian, and L. Washburn. 2017. Persistent spatial structuring of coastal ocean acidification in the California Current System. Scientific Reports, doi: 10.1038/s41598-017-02777-y

Swezey D.S., J.R. Bean, A.T. Ninokawa, T.M. Hill, B. Gaylord, and E. Sanford. 2017. Interactive effects of temperature, food, and skeletal mineralogy mediate biological responses to ocean acidification. Proceedings of the Royal Society B, doi: 10.1098/rspb.2016.2349.

Evans, T.G., M.H. Pespeni, G.E. Hofmann, S.R. Palumbi, and E. Sanford. 2017. Transcriptomic responses to seawater acidification among sea urchin populations inhabiting a natural pH mosaic. Molecular Ecology 26: 2257–2275. doi: 10.1111/mec.14038.

Bible, J.M, K.R. Griffith, and E. Sanford. 2017. Inducible defenses in Olympia oysters in response to an invasive predator. Oecologia, doi: 10.1007/s00442-017-3811-x

Sunday, J.M., K.E. Fabricius, K.J. Kroeker, K.M. Anderson, N.E. Brown, J.P. Barry, S.D. Connell, S. Dupont, B. Gaylord, J.M. Hall-Spencer, T. Klinger, M. Milazzo, P.L. Munday, B.D. Russell, E. Sanford, V. Thiyagarajan, M.L.H. Vaughan, S. Widdicombe, and C.D.G. Harley. 2016. Ocean acidification can mediate biodiversity shifts by changing biogenic habitat. Nature Climate Change 7: 81–85.

Feely, R.A., S. Alin, B. Carter, N. Bednaršek, B. Hales, F. Chan, T.M. Hill, B. Gaylord, E. Sanford, R.H. Byrne, C.L. Sabine, D. Greeley, and L. Juranek. 2016. Chemical and biological impacts of ocean acidification along the West Coast of North America. Estuarine, Coastal and Shelf Science 183: 260–270.

Jellison, B.M., A.T. Ninokawa, T.M. Hill, E. Sanford, and B. Gaylord. 2016. Ocean acidification alters the response of intertidal snails to a key sea star predator. Proceedings of the Royal Society B, doi: 10.1098/rspb.2016.0890

Pfister, C., K. Roy, J.T. Wootton, S. McCoy, R.T. Paine, T.H. Suchanek, and E. Sanford. 2016. Historical baselines and the future of shell calcification for a foundation species in a changing ocean. Proceedings of the Royal Society B, doi: 10.1098/rspb.2016.0392

Kroeker, K.J., E. Sanford, J.M. Rose, C.A. Blanchette, F. Chan, F.P. Chavez, B. Gaylord, B. Helmuth, T.M. Hill, G.E. Hofmann, M.A. McManus, B.A. Menge, K.J. Nielsen, P.T. Raimondi, A.D. Russell, and L. Washburn. 2016. Interacting environmental mosaics drive geographic variation in mussel performance and predation vulnerability. Ecology Letters, doi: 10.1111/ele.12613

Bible, J.M. and E. Sanford. 2016. Local adaptation in an estuarine foundation species: implications for restoration. Biological Conservation 193: 95–102.

Kwan, C.K., E. Sanford, and J. Long. 2015. Copper pollution increases the relative importance of predation risk in an aquatic food web. PLoS ONE 10(7): e0133329. doi:10.1371/journal.pone.0133329.

Evans, T.G., J.L. Padilla-Gamiño, M.W. Kelly, M.H. Pespeni, F. Chan, B.A. Menge, B. Gaylord, T.M. Hill, A.D. Russell, S.R. Palumbi, E. Sanford, and G.E. Hofmann. 2015. Ocean acidification research in the 'post-genomic' era: Roadmaps from the purple sea urchin Strongylocentrotus purpuratus. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology. doi: 10.1016/j.cbpa.2015.03.007

Gaylord, B., K.J. Kroeker, J.M. Sunday, K.M. Anderson, J.P. Barry, N.E. Brown, S.D. Connell, S. Dupont, K.E. Fabricius, J.M. Hall-Spencer, T. Klinger, M. Milazzo, P.L. Munday, B.D. Russell, E. Sanford, S.J. Schreiber, V. Thiyagarajan, M.L.H. Vaughan, S. Widdicombe, C.D.G. Harley.  2014.  Ocean acidification through the lens of ecological theory. Ecology 96: 3–15.

Kroeker, K.J., B. Gaylord, T.M. Hill, J.D. Hosfelt, S.H. Miller, and E. Sanford. 2014. The role of temperature in determining species’ vulnerability to ocean acidification: a case study using Mytilus galloprovincialis. PLOS ONE 9(7): e100353. doi:10.1371/journal.pone.0100353

Kroeker, K.J., E. Sanford, B.M. Jellison, and B. Gaylord. 2014. Predicting the effects of ocean acidification on predator-prey interactions: A conceptual framework based on coastal molluscs.  Biological Bulletin 226: 211–222.

Sanford, E., B. Gaylord, A. Hettinger, E.A. Lenz, K. Meyer, and T.M. Hill. 2014. Ocean acidification increases the vulnerability of native oysters to predation by invasive snails. Proceedings of the Royal Society of London: Biological Sciences Series B 281: 20132681

Hofmann, G. E., T. G. Evans, M. W. Kelly, J. L. Padilla-Gamiño, C. A. Blanchette, L. Washburn, F. Chan, M. A. McManus, B. A. Menge, B. Gaylord, T. M. Hill, E. Sanford, M. LaVigne, J. M. Rose, L. Kapsenberg, and J. M. Dutton. 2014.  Exploring local adaptation and the ocean acidification seascape – studies in the California Current Large Marine Ecosystem. Biogeosciences 11: 1053–1064.

Sanford, E.  2013.  The Biogeography of Marine Communities.  Ch. 7 in: M.D. Bertness, J. Bruno, B.R. Silliman, and J.J. Stachowicz (eds), Marine Community Ecology and Conservation. Sinauer Press, Sunderland, MA.

Hettinger, A., E. Sanford, T. M. Hill, J. D. Hosfelt, A. D. Russell, and B. Gaylord. 2013. The influence of food supply on the response of Olympia oyster larvae to ocean acidification. Biogeosciences 10: 6629–6638.

Howard, J., et al. 2013. Oceans and marine resources in a changing climate. Oceanography and Marine Biology: An Annual Review 51: 71–192.

Hettinger, A., E. Sanford, T.M. Hill, E.A. Lenz, A.D. Russell, and B. Gaylord. 2013. Larval carry-over effects from ocean acidification persist in the natural environment.  Global Change Biology 19: 3317–3326.

Pincebourde, S., E. Sanford, and B. Helmuth.  2013. Survival and arm abscission are linked to regional heterothermy in an intertidal sea star. Journal of Experimental Biology 216: 2183–2191.

Kelly, M.W., R.K. Grosberg, and E. Sanford. 2013. Trade-offs, geography, and limits to thermal adaptation in a tide pool copepod.  American Naturalist 181: 846–854.

Kuo, E.S.L., and E. Sanford. 2013. Distribution of the seaweed limpet Lottia insessa along the Pacific coast.  Pacific Science 67: 303–313.

LaVigne, M. T.M. Hill, E. Sanford, B. Gaylord, A.D. Russell, E.A. Lenz, J.D. Hosfelt, M.K. Young.  2013.  The elemental composition of purple sea urchin (Strongylocentrotus purpuratus) calcite and potential effects of pCO2 during early life stages.  Biogeosciences 10: 3465–3477.

Pespeni, M.H., E. Sanford, B. Gaylord,  T. M. Hill,  J. D. Hosfelt,  H. Jaris, M. LaVigne, E. A. Lenz,  A. D. Russell, M. K. Young,  S. R. Palumbi.  2013.  Evolutionary change during experimental ocean acidification.  Proceedings National Academy of Sciences, doi: 10.1073/pnas.1220673110.

Menge, B.A., and E. Sanford. 2013.  Ecological role of sea stars from populations to meta-ecosystems. Ch. 7 in: J. M. Lawrence (ed.), Starfish: Biology and Ecology of the Asteroidea. Johns Hopkins University Press, Baltimore, MD. 

Hettinger, A., E. Sanford, T.M. Hill, A.D. Russell, K.N. Sato, J. Hoey, M. Forsch, H.N. Page, and B. Gaylord.  2012.  Persistent carry-over effects of planktonic exposure to ocean acidification in the Olympia oyster. Ecology 93: 2758–2768.

Kelly, M.W., R.K. Grosberg, and E. Sanford.  2012.  Love the one you’re with: proximity determines paternity success in the barnacle Tetraclita rubescensMolecular Ecology, DOI: 10.1111/mec.12009.

Pincebourde, S., E. Sanford, J. Casas, and B. Helmuth. 2012. Temporal coincidence of environmental stress events modulates predation rates. Ecology Letters 15: 680–688.

Kelly, M.W., E. Sanford, and R.K. Grosberg.  2012.  Limited potential for adaptation to climate change in a broadly-distributed marine crustacean. Proceedings of the Royal Society of London: Biological Sciences Series B 279: 349–356.  

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.

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.

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