Abstract
Theory and empirical work suggest that coral reefs may exhibit alternative stable states of coral versus macroalgal dominance. However, it is unclear how dispersal of coral and macroalgae among reefs might impact this bistability and the resilience of the coral-dominated state. We develop a mathematical model to investigate how reef cover dynamics are affected by (1) coral and macroalgal dispersal between two reefs and (2) heterogeneous grazer abundances. We find that at low coral and macroalgal dispersal levels, a new type of stable state emerges with both coral and macroalgae present. Furthermore, we show that a reef abundant with coral and grazers can support a coral-dominated stable state in a second reef depauperate of grazers by dispersal of coral larvae. These results help explain previous empirical findings on reefs once thought to be incongruent with traditional coral-macroalgal alternative stable states theory—such as intermediate coral and macroalgal cover stable states and high coral-low grazing scenarios. Our findings indicate that changing dispersal levels (e.g., due to climate change, reef degradation) between reefs changes the possible stable states and the grazing rate at which the coral-dominated state is predicted to be stable. This work demonstrates the relevance of accounting for the level of dispersal among coral reefs or other bistable ecosystems when designing conservation management plans.
This is a preview of subscription content, access via your institution.




References
Amarasekare P (1998) Interactions between local dynamics and dispersal: insights from single species models. Theor Popul Biol 53:44–59. https://doi.org/10.1006/tpbi.1997.1340
Aronson RB, Precht WF (2006) Conservation, precaution, and Caribbean reefs. Coral Reefs 25:441–450. https://doi.org/10.1007/s00338-006-0122-9
Balkenhol N, Cushman SA, Storfer AT, Waits LP (2015) Introduction to landscape genetics–concepts, methods, applications. In: Balkenhol N, Cushman SA, Storfer AT, Waits LP (eds) Landscape Genetics. Wiley, Hoboken, NJ, pp 1–8
Bailey S (2007) Increasing connectivity in fragmented landscapes: an investigation of evidence for biodiversity gain in woodlands. Forest Ecol Manag 238:7–23. https://doi.org/10.1016/j.foreco.2006.09.049
Barnett LA, Baskett ML (2015) Marine reserves can enhance ecological resilience. Ecol Lett 18:1301–1310. https://doi.org/10.1111/ele.12524
Baskett ML, Nisbet RM, Kappel CV, Mumby PJ, Gaines SD (2010) Conservation management approaches to protecting the capacity for corals to respond to climate change: a theoretical comparison. Glob Change Biol 16:1229–1246. https://doi.org/10.1111/j.1365-2486.2009.02062.x
Baskett ML, Fabina NS, Gross K (2014) Response diversity can increase ecological resilience to disturbance in coral reefs. Am Nat 184:E16–E31. https://doi.org/10.1086/676643
Baums IB, Paris CB, Chérubin LM (2006) A bio-oceanographic filter to larval dispersal in a reef-building coral. Limnol Oceanogr 51:1969–1981. https://doi.org/10.4319/lo.2006.51.5.1969
Bellwood DR, Hughes TP, Folke C, Nyström M (2004) Confronting the coral reef crisis. Nature 429:827–833. https://doi.org/10.1038/nature02691
Belsky AJ (1986) Population and community processes in a mosaic grassland in the Serengeti, Tanzania. J Ecol 74:841–856. https://doi.org/10.2307/2260402
Beyer HL, Kennedy EV, Beger M, Chen CA, Cinner JE, Darling ES et al (2018) Risk-sensitive planning for conserving coral reefs under rapid climate change. Conserv Lett 11:e12587. https://doi.org/10.1111/conl.12587
Birrell CL, McCook LJ, Willis BL (2005) Effects of algal turfs and sediment on coral settlement. Mar Pollut Bull 51:408–414. https://doi.org/10.1016/j.marpolbul.2004.10.022
Birrell CL, McCook LJ, Willis BL, Diaz-Pulido GA (2008) Effects of benthic algae on the replenishment of corals and the implications for the resilience of coral reefs. Oceanogr Mar Biol Annu Rev 46:25–63
Blackwood JC, Hastings A, Mumby PJ (2012) The effect of fishing on hysteresis in Caribbean coral reefs. Theor Ecol 5:105–114. https://doi.org/10.1007/s12080-010-0102-0
Blackwood JC, Okasaki C, Archer A, Matt EW, Sherman E, Montovan K (2018) Modeling alternative stable states in Caribbean coral reefs. Nat Resour Model 31. https://doi.org/10.1111/nrm.12157
Bruno JF, Sweatman H, Precht WF, Selig ER, Schutte VG (2009) Assessing evidence of phase shifts from coral to macroalgal dominance on coral reefs. Ecology 90:1478–1484. https://doi.org/10.1890/08-1781.1
Carpenter S, Walker B, Anderies JM, Abel N (2001) From metaphor to measurement: resilience of what to what? Ecosystems 4:765–781. https://doi.org/10.1007/s10021-001-0045-9
Carpenter RC, Edmunds PJ (2006) Local and regional scale recovery of Diadema promotes recruitment of scleractinian corals. Ecol Lett 9:271–280. https://doi.org/10.1111/j.1461-0248.2005.00866.x
Celis-Plá PS, Hall-Spencer JM, Horta PA, Milazzo M, Korbee N, Cornwall CE, Figueroa FL (2015) Macroalgal responses to ocean acidification depend on nutrient and light levels. Front Mar Sci 2:26. https://doi.org/10.3389/fmars.2015.00026
Chauvin A, Denis V, Cuet P (2011) Is the response of coral calcification to seawater acidification related to nutrient loading? Coral Reefs 30:911. https://doi.org/10.1007/s00338-011-0786-7
Cheal AJ, MacNeil MA, Cripps E, Emslie MJ, Jonker M, Schaffelke B, Sweatman H (2010) Coral–macroalgal phase shifts or reef resilience: links with diversity and functional roles of herbivorous fishes on the Great Barrier Reef. Coral Reefs 29:1005–1015. https://doi.org/10.1007/s00338-010-0661-y
Chong-Seng KM, Mannering TD, Pratchett MS, Bellwood DR, Graham NA (2012) The influence of coral reef benthic condition on associated fish assemblages. PLoS ONE 7:e42167. https://doi.org/10.1371/journal.pone.0042167
Côté IM, Darling ES (2010) Rethinking ecosystem resilience in the face of climate change. PLoS Biol 8:e1000438. https://doi.org/10.1371/journal.pbio.1000438
Cowen RK, Sponaugle S (2009) Larval dispersal and marine population connectivity. Annu Rev Mar Sci 1:443–466. https://doi.org/10.1146/annurev.marine.010908.163757
D’Aloia CC, Bogdanowicz SM, Francis RK, Majoris JE, Harrison RG, Buston PM (2015) Patterns, causes, and consequences of marine larval dispersal. PNAS 112:13940–13945. https://doi.org/10.1073/pnas.1513754112
Darling ES, Graham NA, Januchowski-Hartley FA, Nash KL, Pratchett MS, Wilson SK (2017) Relationships between structural complexity, coral traits, and reef fish assemblages. Coral Reefs 36:561–575. https://doi.org/10.1007/s00338-017-1539-z
Deysher L, Norton TA (1981) Dispersal and colonization in Sargassum muticum (Yendo) Fensholt. J Exp Mar Biol Ecol 56:179–195. https://doi.org/10.1016/0022-0981(81)90188-X
Done T (1992a) Phase shifts in coral reef communities and their ecological significance. Hydrobiologia 247:121–132. https://doi.org/10.1007/BF00008211
Done T (1992b) Constancy and change in some great barrier Reef coral communities: 1980–1991. Am Zool 32:655–662. https://doi.org/10.1093/icb/32.6.655
Donovan MK, Friedlander AM, Lecky J, Jouffray JB, Williams GJ, Wedding LM et al (2018) Combining fish and benthic communities into multiple regimes reveals complex reef dynamics. Sci Rep 8:1–11. https://doi.org/10.1038/s41598-018-35057-4
Dudgeon SR, Aronson RB, Bruno JF, Precht WF (2010) Phase shifts and stable states on coral reefs. Mar Ecol Prog Ser 413:201–216. https://doi.org/10.3354/meps08751
Elmhirst T, Connolly SR, Hughes TP (2009) Connectivity, regime shifts and the resilience of coral reefs. Coral Reefs 28:949–957. https://doi.org/10.1007/s00338-009-0530-8
Evensen NR, Doropoulos C, Wong KJ, Mumby PJ (2019) Stage-specific effects of Lobophora on the recruitment success of a reef-building coral. Coral Reefs 38:489–498. https://doi.org/10.1007/s00338-019-01804-w
Fabina NS, Baskett ML, Gross K (2015) The differential effects of increasing frequency and magnitude of extreme events on coral populations. Ecol Appl 25:1534–1545. https://doi.org/10.1890/14-0273.1
Figueiredo J, Baird AH, Harii S, Connolly SR (2014) Increased local retention of reef coral larvae as a result of ocean warming. Nat Clim Change 4:498–502. https://doi.org/10.1038/nclimate2210
Filbee-Dexter K, Scheibling RE (2014) Sea urchin barrens as alternative stable states of collapsed kelp ecosystems. Mar Ecol Prog Ser 495:1–25. https://doi.org/10.3354/meps10573
Fung T, Seymour RM, Johnson CR (2011) Alternative stable states and phase shifts in coral reefs under anthropogenic stress. Ecology 92:967–982. https://doi.org/10.1890/10-0378.1
González-Rivero M, Yakob L, Mumby PJ (2011) The role of sponge competition on coral reef alternative steady states. Ecol Model 222:1847–1853. https://doi.org/10.1016/j.ecolmodel.2011.03.020
Graham NAJ, Nash KL (2013) The importance of structural complexity in coral reef ecosystems. Coral Reefs 32:315–326. https://doi.org/10.1007/s00338-012-0984-y
Graham NAJ, Jennings S, MacNeil MA, Mouillot D, Wilson SK (2015) Predicting climate-driven regime shifts versus rebound potential in coral reefs. Nature 518:94–97. https://doi.org/10.1038/nature14140
Harborne AR, Mumby PJ, Kennedy EV, Ferrari R (2011) Biotic and multi-scale abiotic controls of habitat quality: their effect on coral-reef fishes. Mar Ecol Prog Ser 437:201–214. https://doi.org/10.3354/meps09280
Harrison A, Bjorndal KA (2006) Connectivity and wide-ranging species in the ocean. Conservation Biology Series-Cambridge 14:213
Hock K, Wolff NH, Ortiz JC, Condie SA, Anthony KR, Blackwell PG, Mumby PJ (2017) Connectivity and systemic resilience of the Great Barrier Reef. PLoS Biol 15:e2003355. https://doi.org/10.1371/journal.pbio.2003355
Hoey AS, Bellwood DR (2011) Suppression of herbivory by macroalgal density: a critical feedback on coral reefs? Ecol Lett 14:267–273. https://doi.org/10.1111/j.1461-0248.2010.01581.x
Holling CS (1973) Resilience and stability of ecological systems. Annu Rev Ecol Syst 4:1–24
Hughes TP (1994) Catastrophes, phase shifts, and large-scale degradation of a Caribbean coral reef. Science 265:1547–1551. https://doi.org/10.1126/science.265.5178.1547
Hughes TP (1996) Demographic approaches to community dynamics: a coral reef example. Ecology 77:2256–2260. https://doi.org/10.2307/2265718
Hughes TP, Graham NAJ, Jackson JB, Mumby PJ, Steneck RS (2010) Rising to the challenge of sustaining coral reef resilience. TREE 25:633–642. https://doi.org/10.1016/j.tree.2010.07.011
Hughes TP, Barnes ML, Bellwood DR, Cinner JE, Cumming GS, Jackson JB et al (2017) Coral reefs in the Anthropocene. Nature 546:82–90. https://doi.org/10.1038/nature22901
Ives AR, Carpenter SR (2007) Stability and diversity of ecosystems. Science 317:58–62. https://doi.org/10.1126/science.1133258
Jones GP, Almany GR, Russ GR, Sale PF, Steneck RS, Van Oppen MJH, Willis BL (2009) Larval retention and connectivity among populations of corals and reef fishes: history, advances and challenges. Coral Reefs 28:307–325. https://doi.org/10.1007/s00338-009-0469-9
Jouffray JB, Nyström M, Norström AV, Williams ID, Wedding LM, Kittinger JN, Williams GJ (2015) Identifying multiple coral reef regimes and their drivers across the Hawaiian archipelago. Philos T R Soc B 370:20130268. https://doi.org/10.1098/rstb.2013.0268
Kang Y, Lanchier N (2011) Expansion or extinction: deterministic and stochastic two-patch models with Allee effects. J Math Biol 62:925–973. https://doi.org/10.1007/s00285-010-0359-3
Karatayev VA, Baskett ML (2020) At what spatial scales are alternative stable states relevant in highly interconnected ecosystems? Ecology 101:e02930. https://doi.org/10.1002/ecy.2930
Knipl D, Röst G (2016) Spatially heterogeneous populations with mixed negative and positive local density dependence. Theor Popul Biol 109:6–15. https://doi.org/10.1016/j.tpb.2016.01.001
Knowlton N (1992) Thresholds and multiple stable states in coral reef community dynamics. Am Zool 32:674–682. https://doi.org/10.1093/icb/32.6.674
Koop K, Booth D, Broadbent A, Brodie J, Bucher D, Capone D et al (2001) ENCORE: the effect of nutrient enrichment on coral reefs. Synthesis of results and conclusions. Mar Pollut Bull 42:91–120. https://doi.org/10.1016/S0025-326X(00)00181-8
Krosby M, Tewksbury J, Haddad NM, Hoekstra J (2010) Ecological connectivity for a changing climate. Conserv Biol 24:1686–1689
Kuffner IB, Walters LJ, Becerro MA, Paul VJ, Ritson-Williams R, Beach KS (2006) Inhibition of coral recruitment by macroalgae and cyanobacteria. Mar Ecol Prog Ser 323:07–117. https://doi.org/10.3354/meps323107
Loreau M, Mouquet N, Gonzalez A (2003) Biodiversity as spatial insurance in heterogeneous landscapes. PNAS 100:12765–12770. https://doi.org/10.1073/pnas.2235465100
Manel S, Schwartz MK, Luikart G, Taberlet P (2003) Landscape genetics: combining landscape ecology and population genetics. TREE 18:189–197. https://doi.org/10.1016/S0169-5347(03)00008-9
MATLAB (2019) version 9.6.0 (R2019a). Natick, Massachusetts: The MathWorks Inc
McCauley DJ, Pinsky ML, Palumbi SR, Estes JA, Joyce FH, Warner RR (2015) Marine defaunation: animal loss in the global ocean. Science 347:6219. https://doi.org/10.1126/science.1255641
McCook L, Jompa J, Diaz-Pulido G (2001) Competition between corals and algae on coral reefs: a review of evidence and mechanisms. Coral Reefs 19:400–417. https://doi.org/10.1007/s003380000129
McLaren JR, Jefferies RL (2004) Initiation and maintenance of vegetation mosaics in an Arctic salt marsh. J Ecol 92:648–660. https://doi.org/10.1111/j.0022-0477.2004.00897.x
McManus LC, Watson JR, Vasconcelos VV, Levin SA (2019) Stability and recovery of coral-algae systems: the importance of recruitment seasonality and grazing influence. Theor Ecol 12:61–72. https://doi.org/10.1007/s12080-018-0388-x
McManus LC, Vasconcelos VV, Levin SA, Thompson DM, Kleypas JA, Castruccio FS et al (2020) Extreme temperature events will drive coral decline in the Coral Triangle. Glob Change Biol 26:2120–2133. https://doi.org/10.1111/gcb.14972
Munday PL, Leis JM, Lough JM, Paris CB, Kingsford MJ, Berumen ML, Lambrechts J (2009) Climate change and coral reef connectivity. Coral Reefs 28:379–395. https://doi.org/10.1007/s00338-008-0461-9
Mumby PJ (2006) The impact of exploiting grazers (Scaridae) on the dynamics of Caribbean coral reefs. Ecol Appl 16:747–769. https://doi.org/10.1890/1051-0761(2006)016[0747:TIOEGS]2.0.CO;2
Mumby PJ, Hastings A, Edwards HJ (2007a) Thresholds and the resilience of Caribbean coral reefs. Nature 450:98. https://doi.org/10.1038/nature06252
Mumby PJ, Harborne AR, Williams J, Kappel CV, Brumbaugh DR, Micheli F et al (2007b) Trophic cascade facilitates coral recruitment in a marine reserve. PNAS 104:8362–8367. https://doi.org/10.1073/pnas.0702602104
Mumby PJ (2009) Phase shifts and the stability of macroalgal communities on Caribbean coral reefs. Coral Reefs 28:761–773. https://doi.org/10.1007/s00338-009-0506-8
Mumby PJ, Steneck RS, Hastings A (2013) Evidence for and against the existence of alternate attractors on coral reefs. Oikos 122:481–491. https://doi.org/10.1111/j.1600-0706.2012.00262.x
Nyström M, Folke C, Moberg F (2000) Coral reef disturbance and resilience in a human-dominated environment. TREE 15:413–417. https://doi.org/10.1016/S0169-5347(00)01948-0
O’Connor MI, Bruno JF, Gaines SD, Halpern BS, Lester SE, Kinlan BP, Weiss JM (2007) Temperature control of larval dispersal and the implications for marine ecology, evolution, and conservation. PNAS 104:1266–1271. https://doi.org/10.1073/pnas.0603422104
Paris-Limouzy CB (2011) Reef interconnectivity/larval dispersal. In: Cabioch G, Davies P, Done T, Gischler E, Macintyre IG, Wood R, Woodroffe C (eds) Encyclopedia of modern coral reefs: structure, form and process. Berlin, Germany, pp 881–889
Pauly D, Christensen V, Guénette S, Pitcher TJ, Sumaila UR, Walters CJ et al (2002) Towards sustainability in world fisheries. Nature 418:689–695. https://doi.org/10.1038/nature01017
Petraitis PS, Dudgeon SR (2004) Detection of alternative stable states in marine communities. J Exp Mar Biol Ecol 300:343–371. https://doi.org/10.1016/j.jembe.2003.12.026
Petraitis P (2013) Multiple stable states in natural ecosystems. Oxford, UK
R Core Team (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/
Ratajczak Z, D’Odorico P, Collins SL, Bestelmeyer BT, Isbell FI, Nippert JB (2017) The interactive effects of press/pulse intensity and duration on regime shifts at multiple scales. Ecol Monogr 87:198–218. https://doi.org/10.1002/ecm.1249
Roff G, Mumby PJ (2012) Global disparity in the resilience of coral reefs. TREE 27:404–413. https://doi.org/10.1016/j.tree.2012.04.007
Rogers CS, Miller J (2006) Permanent “phase shifts” or reversible declines in coral cover? Lack of recovery of two coral reefs in St. John, US Virgin Islands. Mar Ecol Prog Ser 306:103–114. https://doi.org/10.3354/meps306103
Scheffer M (1989) Alternative stable states in eutrophic, shallow freshwater systems: a minimal model. Hydrobiol Bull 23:73–83. https://doi.org/10.1007/BF02286429
Scheffer M, Carpenter S, Foley JA, Folke C, Walker B (2001) Catastrophic shifts in ecosystems. Nature 413:591. https://doi.org/10.1038/35098000
Schmitt RJ, Holbrook SJ, Davis SL, Brooks AJ, Adam TC (2019) Experimental support for alternative attractors on coral reefs. PNAS 116:4372–4381. https://doi.org/10.1073/pnas.1812412116
Schröder A, Persson L, De Roos AM (2005) Direct experimental evidence for alternative stable states: a review. Oikos 110:3–19. https://doi.org/10.1111/j.0030-1299.2005.13962.x
Sheppard C, Davy S, Pilling G, Graham N (2017) The biology of coral reefs. Oxford, UK
Srinivasan UT, Cheung WW, Watson R, Sumaila UR (2010) Food security implications of global marine catch losses due to overfishing. J Bioecon 12:183–200. https://doi.org/10.1007/s10818-010-9090-9
Staver AC, Archibald S, Levin SA (2011) The global extent and determinants of savanna and forest as alternative biome states. Science 334:230–232. https://doi.org/10.1126/science.1210465
Steneck RS, Mumby PJ, MacDonald C, Rasher DB, Stoyle G (2018) Attenuating effects of ecosystem management on coral reefs. Sci Adv 4:eaao5493. https://doi.org/10.1126/sciadv.aao5493
Stiger V, Payri CE (1999) Spatial and temporal patterns of settlement of the brown macroalgae Turbinaria ornata and Sargassum mangarevense in a coral reef on Tahiti. Mar Ecol Prog Ser 191:91–100. https://doi.org/10.3354/meps191091
Sumaila UR, Cheung WW, Lam VW, Pauly D, Herrick S (2011) Climate change impacts on the biophysics and economics of world fisheries. Nature Clim Change 1:449–456. https://doi.org/10.1038/nclimate1301
Tebbett SB, Bellwood DR, Purcell SW (2018) Sediment addition drives declines in algal turf yield to herbivorous coral reef fishes: implications for reefs and reef fisheries. Coral Reefs 37:929–937. https://doi.org/10.1007/s00338-018-1718-6
Tebbett SB, Bellwood DR (2019) Algal turf sediments on coral reefs: what’s known and what’s next. Mar Pollut Bull 149:110542. https://doi.org/10.1016/j.marpolbul.2019.110542
Tilman D (1994) Competition and biodiversity in spatially structured habitats. Ecology 75:2–16. https://doi.org/10.2307/1939377
Treml EA, Halpin PN, Urban DL, Pratson LF (2008) Modeling population connectivity by ocean currents, a graph-theoretic approach for marine conservation. Landscape Ecol 23:19–36. https://doi.org/10.1007/s10980-007-9138-y
Treml EA, Roberts JJ, Chao Y, Halpin PN, Possingham HP, Riginos C (2012) Reproductive output and duration of the pelagic larval stage determine seascape-wide connectivity of marine populations. Integr Comp Biol 52:525–537. https://doi.org/10.1093/icb/ics101
van Nes EH, Scheffer M (2005) Implications of spatial heterogeneity for catastrophic regime shifts in ecosystems. Ecology 86:1797–1807. https://doi.org/10.1890/04-0550
Veron JEN (1995) Corals in space and time: the biogeography and evolution of the Scleractinia. Ithaca, USA
Wenger AS, Harris D, Weber S, Vaghi F, Nand Y, Naisilisili W et al (2020) Best-practice forestry management delivers diminishing returns for coral reefs with increased land-clearing. J Appl Ecol 57:2381–2392. https://doi.org/10.1111/1365-2664.13743
Williams I, Polunin N (2001) Large-scale associations between macroalgal cover and grazer biomass on mid-depth reefs in the Caribbean. Coral Reefs 19:358–366. https://doi.org/10.1007/s003380000121
Wilson SK, Bellwood DR, Choat JH, Furnas MJ (2003) Detritus in the epilithic algal matrix and its use by coral reef fishes. Oceanogr Mar Biol 41:279–310
Wilson SK, Graham NA, Pratchett MS, Jones GP, Polunin NV (2006) Multiple disturbances and the global degradation of coral reefs: are reef fishes at risk or resilient? Glob Change Biol 12:2220–2234. https://doi.org/10.1111/j.1365-2486.2006.01252.x
Wood S, Paris CB, Ridgwell A, Hendy EJ (2014) Modelling dispersal and connectivity of broadcast spawning corals at the global scale. Global Ecol Biogeogr 23:1–11. https://doi.org/10.1111/geb.12101
Żychaluk K, Bruno JF, Clancy D, McClanahan TR, Spencer M (2012) Data-driven models for regional coral-reef dynamics. Ecol Lett 15:151–158. https://doi.org/10.1111/j.1461-0248.2011.01720.x
Acknowledgements
We wish to thank Compute Canada for providing computational resources and Z. Ahmed for his guidance in this process. We wish to thank A. McLeod and E. Tekwa for helpful comments on the project.
Funding
This research was supported by the Natural Sciences and Engineering Research Council of Canada (NSERC) Canada Graduate Scholarship to A.G. and NSERC Discovery Grants (no. 4922 to M.K. and no. 5134 to M.-J. F.) and Canada Research Chairs to M.K and M.-J.F.
Author information
Authors and Affiliations
Contributions
All authors contributed to the study conception and design. A.G. led the design of the study, ran the model and wrote the first draft of the manuscript. All authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
Springer Nature or its licensor holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Greiner, A., S. Darling, E., Fortin, MJ. et al. The combined effects of dispersal and herbivores on stable states in coral reefs. Theor Ecol 15, 321–335 (2022). https://doi.org/10.1007/s12080-022-00546-w
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12080-022-00546-w