Coral Reefs

, Volume 36, Issue 4, pp 1291–1305 | Cite as

Large-area imaging reveals biologically driven non-random spatial patterns of corals at a remote reef

  • Clinton B. EdwardsEmail author
  • Yoan Eynaud
  • Gareth J. Williams
  • Nicole E. Pedersen
  • Brian J. Zgliczynski
  • Arthur C. R. Gleason
  • Jennifer E. Smith
  • Stuart A. Sandin


For sessile organisms such as reef-building corals, differences in the degree of dispersion of individuals across a landscape may result from important differences in life-history strategies or may reflect patterns of habitat availability. Descriptions of spatial patterns can thus be useful not only for the identification of key biological and physical mechanisms structuring an ecosystem, but also by providing the data necessary to generate and test ecological theory. Here, we used an in situ imaging technique to create large-area photomosaics of 16 plots at Palmyra Atoll, central Pacific, each covering 100 m2 of benthic habitat. We mapped the location of 44,008 coral colonies and identified each to the lowest taxonomic level possible. Using metrics of spatial dispersion, we tested for departures from spatial randomness. We also used targeted model fitting to explore candidate processes leading to differences in spatial patterns among taxa. Most taxa were clustered and the degree of clustering varied by taxon. A small number of taxa did not significantly depart from randomness and none revealed evidence of spatial uniformity. Importantly, taxa that readily fragment or tolerate stress through partial mortality were more clustered. With little exception, clustering patterns were consistent with models of fragmentation and dispersal limitation. In some taxa, dispersion was linearly related to abundance, suggesting density dependence of spatial patterning. The spatial patterns of stony corals are non-random and reflect fundamental life-history characteristics of the taxa, suggesting that the reef landscape may, in many cases, have important elements of spatial predictability.


Coral reefs Community structure Landscape ecology Spatial dispersion Photomosaics Palmyra Atoll 



This work was made possible through funding provided by the Gordon and Betty Moore Foundation, Grant #3420, and the UC San Diego Frontiers of Innovations Scholarship Program. We are grateful to Gideon Butler, Sho Kodera and Tayler Fewell who contributed to image digitization. This is Palmyra Atoll Research Consortium contribution ## PARC-0125. Thank you to The Nature Conservancy and the Palmyra Atoll Research Consortium for logistical support and the United States Fish Wildlife Service for special use permit # 12533-13025 and access to the refuge.

Supplementary material

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  1. Anderson RM, Gordon DM, Crawley MJ, Hassell MP (1982) Variability in the abundance of animal and plant species. Nature 296:245–248CrossRefGoogle Scholar
  2. Baddeley A, Rubak E, Turner R (2015) Spatial point patterns: methodology and applications with R. Chapman & Hall/CRC Press, Boca RatonGoogle Scholar
  3. Bak R, Engel M (1979) Distribution, abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community. Mar Biol 54:341–352CrossRefGoogle Scholar
  4. Bak R, Termaat R, Dekker R (1982) Complexity of coral interactions: influence of time, location of interaction and epifauna. Mar Biol 69:215–222CrossRefGoogle Scholar
  5. Bellwood DR, Hughes TP, Folke C, Nystrom M (2004) Confronting the coral reef crisis. Nature 429:827–833CrossRefPubMedGoogle Scholar
  6. Bormann FH, Likens G (2012) Pattern and process in a forested ecosystem: disturbance, development and the steady state based on the Hubbard Brook ecosystem study. Springer-Verlag, New YorkGoogle Scholar
  7. Bradbury RR, Young PP (1981) The effects of a major forcing function, wave energy, on a coral reef ecosystem. Mar Ecol Prog Ser 5:229–241CrossRefGoogle Scholar
  8. Bradbury R, Young P (1983) Coral interactions and community structure: an analysis of spatial pattern. Mar Ecol Prog Ser 11:265–271CrossRefGoogle Scholar
  9. Brickner I, Oren U, Frank U, Loya Y (2006) Energy integration between the solitary polyps of the clonal coral Lobophyllia corymbosa. J Exp Biol 209:1690–1695CrossRefPubMedGoogle Scholar
  10. Carlon DB, Olson RR (1993) Larval dispersal distance as an explanation for adult spatial pattern in two Caribbean reef corals. J Exp Mar Bio Ecol 173:247–263CrossRefGoogle Scholar
  11. Chadwick NE (1988) Competition and locomotion in a free-living fungiid coral. J Exp Mar Bio Ecol 123:189–200CrossRefGoogle Scholar
  12. Condit R, Ashton PS, Baker P, Bunyavejchewin S, Gunatilleke S, Gunatilleke N, Hubbell SP, Foster RB, Itoh A, LaFrankie JV, Lee HS, Losos E, Manokaran N, Sukumar R, Yamakura T (2000) Spatial patterns in the distribution of tropical tree species. Science 288:1414–1418CrossRefPubMedGoogle Scholar
  13. Connell JH (1978) Diversity in tropical rain forests and coral reefs. Science 199:1302–1310CrossRefPubMedGoogle Scholar
  14. Connell JH (1985) The consequences of variation in initial settlement vs. post-settlement mortality in rocky intertidal communities. J Exp Mar Bio Ecol 93:11–45CrossRefGoogle Scholar
  15. Dale MRT (1999) Spatial pattern analysis in plant ecology. Ecology 88:366–370Google Scholar
  16. Dale MRT, Dixon P, Fortin M-J, Legendre P, Myers DE, Rosenberg MS (2002) Conceptual and mathematical relationships among methods for spatial analysis. Ecography 25:558–577CrossRefGoogle Scholar
  17. Dana TF (1976) Reef-coral dispersion patterns and environmental variables on a Caribbean coral reef. Bull Mar Sci 26:1–13Google Scholar
  18. Deignan L, Pawlik J (2015) Perilous proximity: does the Janzen-Connell hypothesis explain the distribution of giant barrel sponges on a Florida coral reef? Coral Reefs 34:561–567CrossRefGoogle Scholar
  19. DeVantier L, Endean R (1989) Observations of colony fission following ledge formation in massive reef corals of the genus Porites. Mar Ecol Prog Ser 58:191–195CrossRefGoogle Scholar
  20. Diaz-Pulido G, McCook LJ, Dove S, Berkelmans R, Roff G, Kline DI, Weeks S, Evans RD, Williamson DH, Hoegh-Guldberg O (2009) Doom and boom on a resilient reef: climate change, algal overgrowth and coral recovery. PLoS ONE 4:e5239CrossRefPubMedPubMedCentralGoogle Scholar
  21. Edmunds PJ (2015) A quarter-century demographic analysis of the Caribbean coral, Orbicella annularis, and projections of population size over the next century. Limnol Oceanogr 60:840–855CrossRefGoogle Scholar
  22. Furby KA, Smith JE, Sandin SA (2017) Porites superfusa mortality and recovery from a bleaching event at Palmyra Atoll, USA. Peer J 1:e3204CrossRefGoogle Scholar
  23. Goreau TF (1959) The ecology of Jamaican coral reefs I. Species composition and zonation. Ecology 40:67–90CrossRefGoogle Scholar
  24. Gracias NR, Van Der Zwaan S, Bernardino A, Santos-Victor J (2003) Mosaic-based navigation for autonomous underwater vehicles. IEEE J Oceanic Eng 28:609–624CrossRefGoogle Scholar
  25. Harms KE, Wright SJ, Calderon O, Hernandez A, Herre EA (2000) Pervasive density-dependent recruitment enhances seedling diversity in a tropical forest. Nature 404:493–495CrossRefPubMedGoogle Scholar
  26. Highsmith RC (1982) Reproduction by fragmentation in corals. Mar Ecol Prog Ser 7:207–226CrossRefGoogle Scholar
  27. Highsmith RC, Riggs AC, D’Antonio CM (1980) Survival of hurricane-generated coral fragments and a disturbance model of reef calcification/growth rates. Oecologia 46:322–329CrossRefPubMedGoogle Scholar
  28. Hubbell SP (1979) Tree dispersion, abundance, and diversity in a tropical dry forest. Science 203:1299–1309CrossRefPubMedGoogle Scholar
  29. Hubbell SP, Foster RB (1992) Short-term dynamics of a neotropical forest: why ecological research matters to tropical conservation and management. Oikos 63:48–61CrossRefGoogle Scholar
  30. Hughes R (1987) The functional ecology of clonal animals. Funct Ecol 1:63–69CrossRefGoogle Scholar
  31. Hughes TP (1984) Population dynamics based on individual size rather than age: a general model with a reef coral example. Am Nat 123:778–795CrossRefGoogle Scholar
  32. Hughes T, Jackson J (1980) Do corals lie about their age? Some demographic consequences of partial mortality, fission, and fusion. Science 209:713–715CrossRefPubMedGoogle Scholar
  33. Hughes TP, Connell JH (1987) Population dynamics based on size or age? A reef-coral analysis. Am Nat 129:818–829CrossRefGoogle Scholar
  34. Hughes TP, Tanner JE (2000) Recruitment failure, life histories, and long-term decline of Caribbean corals. Ecology 81:2250–2263CrossRefGoogle Scholar
  35. Hughes TP, Ayre D, Connell JH (1992) The evolutionary ecology of corals. Trends Ecol Evol 7:292–295CrossRefPubMedGoogle Scholar
  36. Hutchinson GE (1953) The concept of pattern in ecology. Proceedings of the Academy of Natural Sciences of Philadelphia 105:1–12Google Scholar
  37. Jackson JBC, Hughes TP (1985) Adaptive strategies of coral-reef invertebrates: coral-reef environments that are regularly disturbed by storms and by predation often favor the very organisms most susceptible to damage by these processes. Am Sci 73:265–274Google Scholar
  38. Jackson JBC, Coates AG (1986) Life cycles and evolution of clonal (modular) animals. Proc R Soc Lond B Biol Sci 313:7–22CrossRefGoogle Scholar
  39. Janzen DH (1970) Herbivores and the number of tree species in tropical forests. Am Nat 104:501–528CrossRefGoogle Scholar
  40. Jolles AE, Sullivan P, Alker AP, Harvell CD (2002) Disease transmission of aspergillosis in sea fans: inferring process from spatial pattern. Ecology 83:2373–2378CrossRefGoogle Scholar
  41. Karlson RH, Cornell HV, Hughes TP (2007) Aggregation influences coral species richness at multiple spatial scales. Ecology 88:170–177CrossRefPubMedGoogle Scholar
  42. Kenyon JC, Maragos JE, Cooper S (2010) Characterization of coral communities at Rose Atoll, American Samoa. Atoll Res Bull 586:1–28CrossRefGoogle Scholar
  43. Kramarsky-Winter E, Loya Y (1996) Regeneration versus budding in fungiid corals: a trade-off. Mar Ecol Prog Ser 134:179–185CrossRefGoogle Scholar
  44. Lewis JB (1970) Spatial distribution and pattern of some Atlantic reef corals. Nature 227:1158–1159CrossRefGoogle Scholar
  45. Lieberman D, Lieberman M, Peralta R, Hartshorn GS (1985) Mortality patterns and stand turnover rates in a wet tropical forest in Costa Rica. J Ecol 73:915–924CrossRefGoogle Scholar
  46. Lirman D, Gracias NR, Gintert BE, Gleason ACR, Reid RP, Negahdaripour S, Kramer P (2007) Development and application of a video-mosaic survey technology to document the status of coral reef communities. Environ Monit Assess 125:59–73CrossRefPubMedGoogle Scholar
  47. Marhaver K, Vermeij M, Rohwer F, Sandin S (2013) Janzen-Connell effects in a broadcast-spawning Caribbean coral: distance-dependent survival of larvae and settlers. Ecology 94:146–160CrossRefPubMedGoogle Scholar
  48. Newman MJH, Paredes GA, Sala E, Jackson JBC (2006) Structure of Caribbean coral reef communities across a large gradient of fish biomass. Ecol Lett 9:1216–1227CrossRefPubMedGoogle Scholar
  49. Pisapia C, Pratchett MS (2014) Spatial variation in background mortality among dominant coral taxa on Australia’s Great Barrier Reef. PLoS ONE 9:e100969CrossRefPubMedPubMedCentralGoogle Scholar
  50. R Core Team (2016) R: a language and environment for statistical computing. R Foundation for Statistical Computing, ViennaGoogle Scholar
  51. Reichelt R, Bradbury R (1984) Spatial patterns in coral reef benthos: multiscale analysis of sites from three oceans. Mar Ecol Prog Ser 17:251–257CrossRefGoogle Scholar
  52. Riegl B, Piller W (2001) Cryptic tissues inside Acropora frameworks (Indonesia): a mechanism to enhance tissue survival in hard times while also increasing framework density. Coral Reefs 20:67–68CrossRefGoogle Scholar
  53. Rietkerk M, van de Koppel J (2008) Regular pattern formation in real ecosystems. Trends Ecol Evol 23:169–175CrossRefPubMedGoogle Scholar
  54. Roff G, Bejarano S, Bozec Y-M, Nugues M, Steneck RS, Mumby PJ (2014) Porites and the phoenix effect: unprecedented recovery after a mass coral bleaching event at Rangiroa Atoll, French Polynesia. Mar Biol 161:1385–1393CrossRefGoogle Scholar
  55. Routledge RD, Swartz TB (1991) Taylor’s power law re-examined. Oikos 60:107–112CrossRefGoogle Scholar
  56. Sandin SA, Smith JE, DeMartini EE, Dinsdale EA, Donner SD, Friedlander AM, Konotchick T, Malay M, Maragos JE, Obura D (2008) Baselines and degradation of coral reefs in the northern Line Islands. PLoS ONE 3:e1548CrossRefPubMedPubMedCentralGoogle Scholar
  57. Sebens KP (1987) The ecology of indeterminate growth in animals. Annu Rev Ecol Evol Syst 18:371–407CrossRefGoogle Scholar
  58. Smith JE, Brainard R, Carter A, Grillo S, Edwards C, Harris J, Lewis L, Obura D, Rohwer F, Sala E, Vroom PS, Sandin S (2016) Re-evaluating the health of coral reef communities: baselines and evidence for human impacts across the central Pacific. Proc R Soc Lond B Biol Sci 283:20151985CrossRefGoogle Scholar
  59. Stimson J (1974) An analysis of the pattern of dispersion of the hermatypic coral Pocillopora meandrina var. nobilis Verril. Ecology: 445–449Google Scholar
  60. Stimson JS (1978) Mode and timing of reproduction in some common hermatypic corals of Hawaii and Enewetak. Mar Biol 48:173–184CrossRefGoogle Scholar
  61. Szmant AM (1986) Reproductive ecology of Caribbean reef corals. Coral Reefs 5:43–53CrossRefGoogle Scholar
  62. Taylor L (1961) Aggregation, variance and the mean. Nature 189:732–735CrossRefGoogle Scholar
  63. Taylor L, Woiwod I (1982) Comparative synoptic dynamics. I. Relationships between inter- and intra-specific spatial and temporal variance/mean population parameters. J Anim Ecol 51:879–906CrossRefGoogle Scholar
  64. Turner MG (1989) Landscape ecology: the effect of pattern on process. Annu Rev Ecol Evol Syst 20:171–197CrossRefGoogle Scholar
  65. Velázquez E, Martínez I, Getzin S, Moloney KA, Wiegand T (2016) An evaluation of the state of spatial point pattern analysis in ecology. Ecography 39:1042–1055CrossRefGoogle Scholar
  66. Wallace CC (1985) Reproduction, recruitment and fragmentation in nine sympatric species of the coral genus Acropora. Mar Biol 88:217–233CrossRefGoogle Scholar
  67. Williams GJ, Smith JE, Conklin EJ, Gove JM, Sala E, Sandin SA (2013) Benthic communities at two remote Pacific coral reefs: effects of reef habitat, depth, and wave energy gradients on spatial patterns. Peer J 1:e81CrossRefPubMedPubMedCentralGoogle Scholar
  68. Wu J (2013) Landscape sustainability science: ecosystem services and human well-being in changing landscapes. Landsc Ecol 28:999–1023CrossRefGoogle Scholar
  69. Zvuloni A, Artzy-Randrup Y, Stone L, Kramarsky-Winter E, Barkan R, Loya Y (2009) Spatio-temporal transmission patterns of black-band disease in a coral community. PLoS ONE 4:e4993CrossRefPubMedPubMedCentralGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany 2017

Authors and Affiliations

  • Clinton B. Edwards
    • 1
    Email author
  • Yoan Eynaud
    • 1
  • Gareth J. Williams
    • 1
    • 2
  • Nicole E. Pedersen
    • 1
  • Brian J. Zgliczynski
    • 1
  • Arthur C. R. Gleason
    • 3
  • Jennifer E. Smith
    • 1
  • Stuart A. Sandin
    • 1
  1. 1.Center for Marine Biodiversity and Conservation, Scripps Institution of OceanographyUniversity of CaliforniaSan Diego, La JollaUSA
  2. 2.School of Ocean SciencesBangor UniversityMenai Bridge, AngleseyUK
  3. 3.Physics DepartmentUniversity of MiamiCoral GablesUSA

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