Land-Sea Interactions and Human Impacts in the Coastal Zone Anson H. Hines ABSTRACT. The Smithsonian Environmental Research Center (SERC) conducts re- search on land-sea interactions to understand natural processes and human impacts in linked ecosystems of the coastal zone. Coastal ecosystems support great biological pro- ductivity and are of immense ecological and economic importance. In addition, more than two-thirds of the human population resides in the coastal zone, where human activities cause chronic and acute disturbance of every habitat and marked degrada- tion of ecological balance and productivity. The Chesapeake Bay and its Rhode River subestuary are used by SERC as model study systems to conduct long-term, intensive monitoring and experiments. Research at SERC focuses on five grand environmental challenges: (I) impacts of atmospheric change on climate ,sea level ,ultraviolet radiation, pollutant deposition ,and carbon balance; (II) impacts of watershed nutrient discharges causing harmful algal blooms, depletion of oxygen, and destruction of submerged veg- etation; (III) food web disruption by pollution and overfishing; (IV) invasive species; and (V) landscape disturbance by agriculture and development. Research by SERC on these grand challenges serves to advise policy and management from improved stewardship of coas traelsources. INTRODUCTION The coastal zone is of immense economic and environmental importance. More than 50% of the Earth’s human population (3 billion people) resides in the coastal zone and relies on the goods and services of coastal ecosystems, and this number is expected to double by 2045 (Creel, 2003). Coastal communities are the most densely populated and fastest growing areas in the United States: 14 of the nation’s largest 20 cities are in coastal locations; more than 50% of the U.S. population lives in 17% of the country’s land, comprising coastal counties; this population concentration is expected increase to 70% within 25 years; and 23 of the 25 most densely populated counties encompass coastal cities and their surrounding sprawl (Crossett et al., 2004). The coastal environment includes the Anso Hn H.ine sS,mithsonia Ennvironment aRle- Earth’s most biologically productive ecosystems, and this diverse environment search Center P, .O B. o x28 6, 47 Contee sWharf includes unmeasured reserves of strategic minerals, oil and gas, and other non- Road E,dgewate rM, arylan d21037-0028 U, SA living resources. The coastal zone encompasses major hubs of global transporta- (hinesa@si.edu M).anuscri pretceiv e 2Ad9ugust tion and commerce and unparalleled opportunities for recreation and tourism, 2008 a;ccepte d2 0Apr i2l009. as well as the majority of fisheries and aquaculture industries. At the same time, 12 e SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES these activities cause chronic and acute disturbance of ev- of Chesapeake Bay. The 585 ha Rhode River subestuary ery coastal habitat: overfishing has removed most large is a shallow (maximum depth = 4 m), soft-bottom em- species at the top of the food web, and coastal waters re- bayment in the lower mesohaline zone of the Bay. The ceive most of the waste of urban centers and agricultural facilities at SERC provide strategic support for research runoff of the coastal plain. at the site and ready access to the rest of the Chesapeake Research at the Smithsonian Environmental Research watershed and estuary. Center (SERC) focuses on land-sea interactions. Scientists at SERC study linked coastal ecosystems to understand natural processes and human impacts in the coastal zone. GRAND CHALLENGES OF COASTAL Ocean productivity is concentrated in the coastal fringe ENVIRONMENTAL RESEARCH where nutrients run off the land and well up from the deep. The coastal environment includes the Earth’s most The purpose of this paper is to present examples that biologically diverse ecosystems: estuaries, wetlands, man- highlight SERC’s coastal research on five grand environ- groves, seagrasses, coral and oyster reefs, kelp forests, and mental challenges. With data sets extending back to the pelagic upwelling areas. Bottom communities and water 1970s and 1980s, SERC research monitors decadal-length column processes of the photic zone are most tightly cou- changes to distinguish seasonal and annual fluctuations pled in the nearshore shallows. Coastal waters comprise from long-term trends in the environment. Importantly, 95% of the oceans’ fisheries. Thus, SERC research focuses SERC research seeks to determine mechanistic under- on improved stewardship of these marine resources. standing of the causes of change at multiple spatial scales ranging from global change to landscape, watershed, eco- system, and community levels of organization. The land CHESAPEAKE BAY AND THE RHODE RIVER and long-term studies at SERC’s Rhode River site afford SUBESTUARY AS A MODEL SYSTEM multidisciplinary experimental analyses of mechanisms controlling ecological interactions. The research there The Smithsonian Environmental Research Center addresses the grand challenges and advises environmen- utilizes the nation’s largest estuary, Chesapeake Bay and tal policy and management for improved stewardship of its 177,000 km? watershed including six states and the coasta lresources. District of Columbia (Figure 1), as its primary research landscape and main study site. In addition to SERC, this GRAND CHALLENGE |: IMPACTS OF ATMOSPHERIC CHANGE study area includes the Smithsonian’s museum complex, zoological exhibits, and administrative offices. An area Human alterations of the atmosphere are causing with a long American history of exploitation of coastal re- rapid changes in climate, sea level, ultraviolet radiation, sources, the Chesapeake watershed is home to 17 million pollutant deposition, and ecosystem carbon balance. Re- people, who are mostly concentrated in the urban centers search by SERC on the salt marshes of the Rhode River and suburban sprawl of Baltimore, Washington, D.C., and subestuary provides a good example of the ecological com- Norfolk. Agriculture, particularly row crops, is the major plexities of this challenge. B. G. Drake and colleagues have land use of the Chesapeake watershed, and farming has been conducting the world’s longest running experimental been the major source of disturbance to the eastern de- manipulation of CO) on natural plant communities (1985 ciduous forest for 400 years. to present), which has been testing the effects of rising at- Established in 1965, SERC owns a unique 1,072 ha mospheric CO, concentration in these salt marshes. The land holding for long-term descriptive and experimental experiment measures response of the two dominant plant studies of linked ecosystems in a model subestuary and species at the site: Spartina patens and Scirpus olneyi. The subwatershed of Chesapeake Bay—the Rhode River, experiment applied nine treatment combinations of three which is located 40 km east of Washington, D.C., and CO, levels in open-top chambers (ambient air at 340 ppm; 10 km south of Annapolis, Maryland (Figure 2). The elevated CO, at a twofold increase in concentration of 680 property at SERC includes cropland, forests in various ppm; and a control treatment without chambers) crossed successional stages, wetlands, and 26 km of undeveloped with types of patches (nearly monospecific S. patens; nearly shoreline; this is the largest contiguous block of land monospecific S. olneyi; and patches with mixes of the two dedicated to environmental research, science education, species) (Drake et al., 1989). Chambers were replaced public access, and stewardship on the western shoreline exactly on replicate marked plots of the nine treatment NUMBER 38 e¢ 13 BCahyesapeake 50 km FIGURE 1 .Map o fChesapeake Bay and its watershed with six physiographic provinces .Arrow indicates the location o fthe Smithsonian Environmenta lResearch Center on the Rhode River subestuary and watershed .Darkened areas indicate 17 clusters o f500 subwatersheds tha tdif- fered in land use and were monitored fo rstream discharges o fnutrients. 14 e SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES [ -R]hod Reive Wr atershed HM SME RpCroperty Privat we it ShER Ccovenant Privat we it heasement P(Priva utenprotected Private Smithsonian Environmental Research Center Bay * 4Os FIGURE 2 .Map of land holdings (shaded green) of the Smithsonian Environmenta lResearch Center (SERC) surrounding the Rhode River subestuary .Red outline show sthe boundar yo fthe watershed. combinations for the duration of the growing season for tion of the plants, which appears to be sequestered in the the past 23 years (1995-2008). Photosynthesis and respi- peat-forming roots of the salt marsh (Carney et al., 2007). ration were measured in each chamber during the growing Research conducted by J. P. Megonigal and colleagues at season, and plant production was measured at the end of the same marsh study site compares effects of increased each season. As predicted, Spartina patens is a C, plant CO, interacting with nutrient additions to the marsh to that responds weakly to rising CO, whereas growth and determine whether peat accumulation is sufficient to keep production were greatly stimulated in Scirpus olneyi as a up with rising sea level. Their initial results indicate that C; plant (Drake and Rasse, 2003). However, the amount the peat accumulation is equivalent to the current rate of stimulation of S. olneyi is significantly inversely depen- of sea-level rise of approximately 3 mm year’, allow- dent on salinity (i.e., water stress), with lower production ing the marsh to persist instead of becoming submerged. in years of high salinities (i.e., low rainfall) (Rasse et al., Additionally, a nonnative species, Phragmites australis, is 2005; and Figure 3). rapidly invading the marsh site, similar to most others Salt marsh research at SERC’s Rhode River site also in the region (King et al., 2007); and its responses to the explores other ecosystem complexities. New research is interaction of rising CO, and nutrients are unknown. The tracking the fate of the carbon added by growth stimula- Chesapeake region has high levels of mercury deposition NUMBER 38 15 (A) (%S)timulation 200 300 400 500 600 700 800 Precipitation (mm) (B) (%S)timulation Salinity (ppt) FIGURE 3. Effect of (A) precipitation and (B) salinity (ppt = parts per thousand) on the stimulation of photosynthesi sb ytwofold increase in CO c, oncentration on the sedge Scirpu solney in open-top chambers placed on a salt marsh o fthe Rhode River subestuary during a 17-year period (1989-2003) .(After Rasse e tal .2, 005.) 16 e SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES that is derived from coal-burning power plants. New work at the SERC salt marsh site shows that microbes rapidly activate the mercury (mercury-methylation) (Mitchell et al., 2008) deposited into marshes, thus feeding it into bio- logical processes on the coastal food web (C. Mitchell and Oo C. Gilmour, Smithsonian Environmental Research Center, 2008, personal communication). a GRAND CHALLENGE II: (mg/NL)itrate-N IMPACTS O FNUTRIENT LOADING o @410 308 @409 Over-enrichment of coastal waters with nutrients causes harmful algal blooms, depletion of oxygen, and ND destruction of submerged vegetation. Eutrophication in Chesapeake Bay and many other coastal systems is caus- — ing “dead zones” of anoxic and hypoxic waters along deeper bottom areas. A major focus of the restoration 0 10 20 30 40 50 60 70 efforts of the Environmental Protection Agency’s Chesa- peake Bay Program has been to reduce nutrient loading Percentage of Cropland by phosphorus and nitrogen runoff into the Bay. Long- term watershed and estuarine water quality monitoring FIGURE 4. Effects of cropland on stream discharge of nitrogen by SERC at the Rhode River site and throughout Chesa- for watersheds in the Piedmont and Coasta lPlain physiographic peake Bay shows the dynamic interactions of stream dis- provinces of Chesapeake Bay (see Figure 1) .Nitrogen is shown as charge, nutrient inputs, and plankton responses affecting nitrate concentration on the y-axis; cropland is shown as a per- oxygen levels. centage of land use of the subwatershed area on the x-axis .(After Watershed nutrient discharge occurs primarily in Jordan et al. ,1997.) storm events and is related to both geologic position (e.g., Piedmont or Coastal Plain provinces of the Chesa- peake watershed) and land use, especially development and agriculture (Figure 4). Plankton productivity is much emphasizes the linkage of submerged aquatic vegetation to higher in years with high runoff, which leads to plank- watershed characteristics (Li et al., 2007). ton blooms (Figure 5). Long-term monitoring from 1986 to 2004 shows that water clarity (Secchi disc depth) and GRAND CHALLENGE III: FooD Wes DISRUPTION near-bottom oxygen levels have declined significantly in BY POLLUTION AND OVERFISHING the Rhode River subestuary (Figure 6). Although oxygen levels at SERC’s long-term monitoring station in the shal- Pollution and overfishing result in severe disruptions low edge of the Bay generally do not fall below alarm- of coastal food webs (Jackson et al., 2001). The combined ing levels of approximately 6 ppm, oxygen levels in the effects of low dissolved oxygen and loss of submerged deeper mainstem of the Bay drop to very low levels (Hagy aquatic vegetation comprise much of the major impact et al., 2004) and occasionally spill into the mouth of the of pollution in coastal systems such as Chesapeake Bay. Rhode River, killing benthic organisms (A. Hines, personal However, inputs of mercury and other toxic chemicals also observations). markedly affect the food web as they become concentrated With the decline in water clarity, light levels are not at its upper levels, often causing serious effects on seafood sufficient to support growth of seagrasses and other sub- that affect human health (Krabbenhoft et al., 2007). Im- merged aquatic vegetation, which had largely disappeared pacts of overfishing and habitat loss have resulted in the from the Rhode River subestuary and much of Chesapeake loss of sustainable stocks for nearly every fishery species in Bay by the early 1970s. These structured ecosystems are Chesapeake Bay and in nearly every coastal system world- important nursery habitats for fish and crabs in coastal wide. After a century of intense exploitation, disease, and systems such as Chesapeake Bay. Recent SERC research ecosystem impacts, oysters, as the Bay’s most productive NUMBER 38 ¢ 17 fishery historically, are now at only 1% of their biomass Non-bloom in 1900 (Rothschild et al., 1994). Eutrophication and year (1999) overfishing act as multiple stressors on coastal food webs, —— Bloom year and management’s too narrow focus on single factors may have adverse consequences for restoring ecosystem health and fishery production (Breitburg et al., 2009). d ’mDP')ra*oi lCdyu c(gtion Blue crabs are the remaining major lucrative fishery in the upper Bay, but the blue crab stock has also declined by 60% since 1991 (CBSAC, 2008). Research by SERC at the Rhode River subestuary provides the most detailed analy- sis of blue crab ecology available (Hines, 2007). Nearly 30 years of SERC experiments show that blue crabs are the dominant predator on benthic communities in the estuary, =. =o ae - > 1 L S l 0 60 120 180 240 300 360 and their foraging limits abundance and species compo- sition of infaunal invertebrates as well as causing major Day of Year bioturbation of the upper 10 cm of sediments (Hines et al., 1990). Long-term monitoring of fish and blue crabs FIGURE 5. Comparison of carbon production in the Rhode River throughout the Rhode River subestuary shows the marked subestuary during two years, one with and one without a spring seasonal and annual variations in population abundance plankton bloom ,which is mainly regulated by variation in spring (Figure 7), as blue crabs migrate from the nursery habitat precipitation and watershed discharge. (After Gallegos and Jor- and become inactive below 9°C in winter. Annual varia- dan 1,997.) tion in recruitment into the Rhode River causes more than a 10-fold fluctuation in abundance, with obvious variation in effects of predation on infaunal invertebrates. Many up- per Chesapeake Bay nursery habitats now appear to be be- low carrying capacity for juvenile blue crabs (Hines et al., 2008). Recent SERC blue crab research has focused on de- 2.0 a Rhod eRiver M, aryland (m D)F eSepebtcrhcuhairy m”B )( og Dlat.tOyoe.mr December-Ma arvcehrage 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 Year FIGURE 6 .Long-term trends in water clarity as determined by Secch i(disk) depth (left) and in oxygen concentration (D.O .= dissolved oxygen; right )in the Rhode Rive rsubestuary .(Figure courtesy o fC .Gallegos.) 18 e© SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 250 200 150 tra p cwmM erblar oelnbunause ntomhflbyer 100 50 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 Year FIGURE 7 .Seasona land annua lvariation in abundance of blue crabs caught in 3 m otter trawls in the Rhode River subestuary .Abundance is the monthly mean o fthree trawls a teach o fthree permanen tstations within t hesetuary. veloping innovative approaches to restoring the blue crab sions (Figure 8). Invasions are dynamic and ongoing in population in the Bay, especially by testing the feasibility Chesapeake Bay, as indicated by recent records of Chi- of releasing hatchery-reared juvenile blue crabs into nurs- nese mitten crabs (Ruiz et al., 2006). Many species are ery areas such as the Rhode River (Hines et al., 2008). having large but poorly understood impacts in Chesa- peake ecosystems, such as the salt marsh reed Phragmites GRAND CHALLENGE IV: INVASIVE SPECIES australis (King et al., 2007). Invasions of nonindigenous species are drastically alter- GRAND CHALLENGE V: ing biodiversity, structure, and function of coastal ecosys- LANDSCAP EDISTURBANC EB YAGRICULTURE tems (Ruiz et al., 2000). The largest, most comprehensive AN DEVELOPMENT research program on marine invasive species in the USA is conducted by SERC. Rates of invasion into coastal Agriculture and urbanization are causing widespread ecosystems are increasing markedly as a result of a wide modifications of landscape structure. Researchers at SERC range of human-mediated vectors, but most importantly recently analyzed various indicators of estuarine habitat as a result of shipping, both ballast water discharge and quality for 31 Chesapeake subwatersheds that differed in hull fouling (Ruiz et al., 2000). The SERC database for five categories of land use composition: forest, agriculture, invasive species (NEMESIS) documents more than 500 developed, mixed agriculture, and mixed developed (Fig- invasive species of invertebrates, algae, and fish in North ure 9). These land uses have profound effects on estuarine American coastal waters. For Chesapeake Bay approxi- habitat quality because they increase stormwater runoff and mately 176 species are documented as established inva- loading of nutrients. Nitrogen discharge into subestuaries of NUMBER 38 e¢ 19 Taxonomic groups of NIS introduced and established in the Chesapeake Bay region (n= 176) Other Vertebrates @ Regular Residents Fishes @ Boundary Residents Invertebrates Vascular Plants Algae 40 60 80 Number of species FIGURE 8 .Number so finvasive specie sdocumented fo ralgae ,vascula rplants ,invertebrates ,fishes ,and other vertebrates (tota lnumber = 176 species )in Chesapeake Bay .Regular residents are species living in habitats below tida linfluence ;boundary residents are species primarily living eithe rabove the intertida lzone o rin non- tida lfreshwater and tha toccasionally move into tida lportions o fthe Bay .(NIS = noninvasive species.) agricultural and developed watersheds was high in both wet CONCLUSION and dry years, but in dry years it was high only in developed watersheds, which continue to have high human water use The decadal data sets generated by SERC for the regardless of rainfall (Figure 10) (Brooks et al., 2006). Land linked ecosystems of the Rhode River and Chesapeake Bay use also has marked effects on levels of toxic chemicals in clearly show the importance of sustaining long-term, in- the food webs of the subestuaries. Level of polychlorinated tensive studies to distinguish natural variation and trends biphenyls (PCBs) was highly correlated with percentage of of human impacts. The rate of change associated with hu- developed lands on the subwatershed (Figure 11). man impacts is increasing markedly as the effects of global In addition to effects on the watershed, development change become manifest and as the human population of of the shoreline has large impacts on coastal ecosystems. the watershed continues to grow rapidly, with another Research by SERC in the Rhode River shows that the 50% increase predicted in the next 25 to 50 years. The shallowest fringe of the subestuary serves as a critical ref- interactive effects of these multiple stressors require much uge habitat for juvenile fishes and crabs to avoid larger more research to define improved management solutions predators, which are restricted to deeper water (Ruiz et to restore and sustain these resources. Scientists at SERC al., 1993; Hines and Ruiz, 1995). Coarse woody debris also extend studies of the large-scale systems of the Rhode from forested shores also plays a valuable role as struc- River and Chesapeake Bay through comparative studies tural habitat and refuge from predators (Everett and with other coastal areas, especially latitudinal compari- Ruiz, 1993). As development results in cutting down the sons of systems in the Smithsonian Marine Science Net- riparian forest and hardening the shoreline with bulk- work along the western Atlantic. Although each site has heads and riprap to prevent erosion, water depth at the its idiosyncratic traits, the common impacts of the grand shoreline increases and the source of woody debris is challenges of atmospheric change, nutrient loading, food lost. With the loss of functional refuge in the nearshore web disruption by pollution and overfishing, invasive spe- shallows, juvenile fish and crabs become increasingly ac- cies, and land development are all manifested pervasively cessible to predators. in the linked ecosystems throughout the coastal zone. 20 * SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Jones Falls Gwynns Falls Patapsco eu angford Curt . BZ \ Southeast MSaegvoetnh : ss Corsica out Rhe ian Battle ade Avon “7 St. Clements 4 A" si t: i lLeonard Wicomico Bis oni “4 Tisees > Ne" Manokin Nomini Totuskey LANDUSE — Agricultural ard Developed eal Forested Mixed-Ag ened Mixed-Dev Elizabeth a ees Kilometers G12'5'25 50 75 100 FIGURE 9 .Map of 31 subwatersheds of Chesapeake Bay that were sampled for effects of land us eon estuarin ehabitats W. atershed swer ecategorized in th efiv epredominan tcategorie shown: fores at,gricultur ed,evelope dm, ixed-agricultur ea,n md ixed-developed. NUMBER 38 21 2500 N(2O)Ooo (pg/LS)u Nrfa cTeo-twaal ter 1500 Forested Mixed-Dev Developed Forested Mixed-Dev Developed Mixed-Ag Agricultural Mixed-Ag Agricultural Watershed Land-use Class FIGURE 10 .Effec to fland use on nitrogen discharge from watersheds in the five land use categories shown in Figure 9 .Stream surface discharges are compared among land use categories between a dry yea rwith record low rainfall (2002, left) and a wet year (2003, right) with high rainfall. (After Brooks et al., 2006.) y = 19.781x - 8.9308 R’= 0.99 W P(pehpr i ictPbnTehC)oBtasl Lyi 0) 10 20 30 40 IDW (d-‘) Percent Commercial Land in Watershed FIGURE 11. Concentration of toxic polychlorinated biphenyls (PCBs) in white perc h(Moron eamericana s)ample dfrom Chesapeak esubestuarie swit hwatersheds o fvarying percentage so fcommercially developed land use (IDW = inverse distance weighted) .Watersheds sampled are shown in Figure 9 .(After King et al. ,2004.) PA POA SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ACKNOWLEDGMENTS Everett ,R .A. ,and G .M .Ruiz .1993 .Coarse Woody Debris as a Refuge from Predation in Aquati cCommunities A. n Experimenta Tl est. Oecolog (iaBerlin 9)3, :475-486. I thank my colleagues at SERC for use of their data Gallegos ,C .L. ,and T .E .Jordan .1997 .Seasona lProgression of Fac- and publications to illustrate this paper, especially De- tor sLimiting Phytoplankton Pigmen tBiomas sin th eRhod eRiver nise Breitburg, Bert Drake, Chuck Gallegos, Cindy Estuary M, aryland (USA) I. C. ontrol son Phytoplankton Growth.Marin Eecolog Pyrogres Sserie s1,61:185-198. Gilmour, Tom Jordan, Pat Megonigal, Rick Osman, Hagy ,J .D. ,W .R .Boynton ,C .W .Keefe ,and K .V .Wood .2004 .Hypoxia and Greg Ruiz. I thank the Smithsonian Marine Science in Chesapeake Bay 1, 950-2001 L: ong-Term Change in Relation to Network and the Smithsonian Environmental Sciences Nutrien tLoading and Rive rFlow E. stuaries 2, 7:634-658. Program for long-term funding to SERC’s Rhode River Hines ,A .H .2007 .Ecology o fJuvenile and Adul tBlue Crabs .In Biologyo fthe Blue Crab ,V .S .Kennedy and L .E .Cronin ,eds. ,pp .565-654. research program. Thanks go to Michael Lang for his Colleg ePark M: arylan dSe aGran Ct olleg eProgram. steady assistance in managing the Marine Science Net- Hines, A. H., A. M. Haddon, and L. A. Wiechert. 1990. Guild Struc- work and for organizing the symposium and editing ture and Foraging Impac to fBlue Crabs and Epibenthic Fish in aSubestuar yo Cf hesapeak eBay M. arin eEcolog yProgres sSeries, these proceedings. 67:105-126. Hines, A. H., E. G. Johnson, A. C. Young, R. Aguilar, M. A. Kramer, M .Goodison ,O .Zmora ,and Y .Zohar .2008 .Release Strategies fo rEstuarine Specie swith Complex Migratory Life Cycles :Stock Enhancemen ot Cf hesapeak eBlu eCrabs C, allinecte sapidus R. e- LITERATURE CITED view si nFisherie sScience 1,6:175-185.Hines ,A .H. ,and G .M .Ruiz .1995 .Tempora lVariation in Juvenile Blue Crab Mortality :Nearshore Shallow sand Cannibalism in Chesa- Breitburg, D. L., J. K. Craig, R. S. Fulford, K. A. Rose, W. R. Boyn- peak eBay B. ulletin o Mf arin eScience 5, 7:885-902. ton, D. Brady, B. J. Ciotti, R. J. Diaz, K. D. Friedland, J. D. Hagy Jackson, J. B. C., M. X. Kirby, W. H. Berger, K. A. Bjorndal, L. W. Bots- Ill, D.R. Hart, A. H. Hines, E. D. Houde, S. E. Kolesar, S. W. ford ,B. J. Bourque ,R. H. Bradbury ,R. Cooke ,J. Erlandson ,J. A. Nixon, J. A. Rice, D. H. Secor, and T. E. Targett. 2009. Nutrient Estes, T. P. Hughes, S. Kidwell, C. B. Lange, H. S. Lenihan, J. M. Enrichmen tand Fisherie sExploitation I:nteractiv eEffect son Es- Pandolfi, C. H. 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NUMBER 38 e¢ 23 Ruiz ,G. ,L .Fegley ,P .Fofonoff ,Y .Cheng ,and R .Lemaitre .2006 .First America P: attern sand Processes A. nnua Rl eview sin Ecolog yand Record so fEriochei rsinensi sH M. ilne Edwards 1, 85 3(Crustacea: Systemat 3ic1s:,481-531. Brachyura V: arunidae )fo rChesapeak eBa yand th eMid-Atlantic Ruiz, G. M., A. H. Hines, and M. H. Posey. 1993. Shallow Water as a Coas to fNorth America A. quati cInvasions 1, :137-142. Refug eHabita fto rFis han dCrustacean si nNon-vegetate dEstuar- Ruiz, G. M., P. W. Fofonoff, J. T. Carlton, M. Wonham and A. H. ies :An Example from Chesapeake Bay M. arine Ecolog yProgress Hines .2000 .Invasion o fCoasta lMarine Communities o fNorth Seri e9s9,:1-16. Hines, Anson H. 2009. "Land–Sea Interactions and Human Impacts in the Coastal Zone." Proceedings of the Smithsonian Marine Science Symposium 38, 11–23. 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