DOI: https://doi.org/10.3354/meps14952
copiedSize-specific predation on an overgrazing sea urchin reveals dominant role of large predatory fish
ABSTRACT:
Predators can play a key role in structuring ecological communities through top-down control, which may be influenced by size-specific relationships, abundance and richness of predatory guilds. In southeastern Australia, the diadematid sea urchin Centrostephanus rodgersii reaches high abundances, overgrazing kelp and maintaining extensive barrens. Despite its ecological importance, predator identity and size-specific nature of predation on this urchin remain incomplete. Here, we conducted urchin tethering assays with remote underwater videos to identify predators across a full size range of urchins (16-110 mm test diameter). We identified 5 predatory fishes from a total of 94 arrays and 61 predation events from 7 sites in the Sydney region, Australia. The eastern blue groper Achoerodus viridis (Labridae) accounted for 75% of events, and the other 4 predatory fishes accounted for the remainder, which involved small urchins only (≤25 mm test diameter). Overall, predation risk decreased as urchin size increased, while time to predation decreased with predator richness. Smaller urchins were more likely attacked from the aboral side, while larger urchins were flipped and attacked from the oral surface. A. viridis was the only predator observed consuming all urchin sizes. These findings show that predator size and richness are key factors determining the predation pressure on C. rodgersii. As many of these factors have been heavily impacted by historical overfishing, regulating urchin populations through effective predator management may be crucial for sustaining ecosystem services in kelp forests.
KEYWORDS
- Video (m770p083_VS1.mp4)
Video S1. Footage of all urchin predators captured throughout the tethering experiment.
- Video (m770p083_VS2.mp4)
Video S2. A female blue groper can be seen foraging around and completely ignoring the readily available urchin.
- Video (m770p083_VS3.mp4)
Video S3. A male blue groper maneuvers carefully to avoid longspined sea urchins while attacking the test by flipping the urchin over and going straight to an oral attack on the urchin's mouth.
Allen GR, Midgley SH, Allen M (2002) Field guide to the freshwater fishes of Australia. Western Australian Museum, Perth
Andrew NL, Byrne M (2001) The ecology of Centrostephanus rodgersii. Dev Aquacult Fish Sci 32:149-160
Andrew NL, O’Neill AL (2000) Large-scale patterns in habitat structure on subtidal rocky reefs in New South Wales. Mar Freshw Res 51:255-263 Crossref
Andrew NL, Underwood AJ (1989) Patterns of abundance of the sea urchin Centrostephanus rodgersii (Agassiz) on the central coast of New South Wales, Australia. J Exp Mar Biol Ecol 131:61-80 Crossref
Andrew NL, Underwood AJ (1993) Density-dependent foraging in the sea urchin Centrostephanus rodgersii on shallow subtidal reefs in New South Wales, Australia. Mar Ecol Prog Ser 99:89-98 Crossref
Andrialovanirina N, Ponton D, Behivoke F, Mahafina J, Léopold M (2020) A powerful method for measuring fish size of small-scale fishery catches using ImageJ. Fish Res 223:105425 Crossref
Antiqueira PAP, Petchey OL, Rezende F, Machado Velho LF, Rodrigues LC, Romero GQ (2022) Warming and top predator loss drive direct and indirect effects on multiple trophic groups within and across ecosystems. J Anim Ecol 91:428-442 Crossref
Aronson RB, Heck KL Jr (1995) Tethering experiments and hypothesis testing in ecology. Mar Ecol Prog Ser 121:307-309 Crossref
Balemi CA, Shears NT (2023) Emergence of the subtropical sea urchin Centrostephanus rodgersii as a threat to kelp forest ecosystems in northern New Zealand. Front Mar Sci 10:1224067 Crossref
Banks SC, Piggott MP, Williamson JE, Bové U, Holbrook NJ, Beheregaray LB (2007) Oceanic variability and coastal topography shape genetic structure in a long-dispersing sea urchin. Ecology 88:3055-3064 PubMed Crossref
Bass NC, Day J, Guttridge TL, Mourier J, Knott NA, Vila Pouca C, Brown C (2021) Residency and movement patterns of adult Port Jackson sharks (Heterodontus portusjacksoni) at a breeding aggregation site. J Fish Biol 99:1455-1466 PubMed Crossref
Bell JD, Burchmore JJ, Pollard DA (1978) Feeding ecology of three sympatric species of leatherjackets (Pisces: Monacanthidae) from a Posidonia seagrass habitat in New South Wales. Mar Freshw Res 29:631-643 Crossref
Bennett S, Wernberg T, Connell SD, Hobday AJ, Johnson CR, Poloczanska ES (2016) The ’Great Southern Reef’: social, ecological and economic value of Australia’s neglected kelp forests. Mar Freshw Res 67:47-56 Crossref
Boada J, Arthur R, Farina S, Santana Y, Mascaró O, Romero J, Alcoverro T (2015) Hotspots of predation persist outside marine reserves in the historically fished Mediterranean Sea. Biol Conserv 191:67-74 Crossref
Boada J, Sanmartí N, Selden RL, Lucas A, Pérez M, Alcoverro T, Romero J (2015) Evaluating potential artifacts of tethering techniques to estimate predation on sea urchins. J Exp Mar Biol Ecol 471:17-22 Crossref
Booth DJ (2010) Natural history of Sydney’s marine fishes: where south meets north. In: Lunney D, Hutchings P, Hochuli D (eds) The natural history of Sydney. Royal Zoological Society of New South Wales, Mosman, p 143-153 Crossref
Burt JM, Tinker MT, Okamoto DK, Demes KW, Holmes K, Salomon AK (2018) Sudden collapse of a mesopredator reveals its complementary role in mediating rocky reef regime shifts. Proc R Soc B 285:20180553 PubMed Crossref
Byrne M, Andrew NL (2020) Centrostephanus rodgersii and Centrostephanus tenuispinus. In: Lawrence JM (ed) Sea urchins: biology and ecology. Elsevier, London, p 379-396 Crossref
Caffo B (2015) Package ’exactLoglinTest’. Link
Cappo M, Harvey E, Malcolm H, Speare P (2003) Potential of video techniques to monitor diversity, abundance and size of fish in studies of marine protected areas. In: Beumer JP, Grant A, Smith DC (eds) Aquatic protected areas—What works best and how do we know? World Congress on Aquatic Protected Areas Proceedings, Cairns, p 455-464
Clemente S, Hernández J, Rodríguez A, Brito A (2010) Identifying keystone predators and the importance of preserving functional diversity in sublittoral rocky-bottom areas. Mar Ecol Prog Ser 413:55-67 Crossref
Clemente S, Hernández JC, Montaño-Moctezuma G, Russell MP, Ebert TA (2013) Predators of juvenile sea urchins and the effect of habitat refuges. Mar Biol 160:579-590 Crossref
Cohen JE, Pimm SL, Yodzis P, Saldaña J (1993) Body sizes of animal predators and animal prey in food webs. J Anim Ecol 62:67-78 Crossref
Coleman MA (2013) Connectivity of the habitat-forming kelp, Ecklonia radiata within and among estuaries and open coast. PLOS ONE 8:e64667 PubMed Crossref
Costa GC (2009) Predator size, prey size, and dietary niche breadth relationships in marine predators. Ecology 90:2014-2019 PubMed Crossref
Cox DR (1972) Regression models and life-tables. J R Stat Soc B 34:187-202 Crossref
Day JK, Knott NA, Swadling DS, Ayre DJ (2021) Dietary analysis and mesocosm feeding trials confirm the eastern rock lobster. Mar Freshw Res 72:1220-1232 Crossref
Day JK, Knott NA, Swadling DS, Huggett MJ, Gaston TF (2024) Assessing lobster and co-predator feeding rates on barrens-forming sea urchins in South East Australia. Front Mar Sci 11:1418506 Crossref
Day JK, Huggett MJ, Gaston TF (2025) Suspected key predators of long-spined urchins fail to show signs of significant predation in gut contents analyses. Estuaries Coasts 48:88 Crossref
Denno RF, Finke DL (2006) Multiple predator interactions and food-web connectance: implications for biological control. In: Brodeur J, Boivin G (eds) Trophic and guild in biological interactions control. Springer, Dordrecht, p 45-70
DiFiore BP, Stier AC (2023) Variation in body size drives spatial and temporal variation in lobster-urchin interaction strength. J Anim Ecol 92:1075-1088 PubMed Crossref
Ebert TA (1982) Longevity, life history, and relative body wall size in sea urchins. Ecol Monogr 52:353-394 Crossref
Eger AM, Marzinelli EM, Beas-Luna R, Blain CO and others (2023) The value of ecosystem services in global marine kelp forests. Nat Commun 14:1894 PubMed Crossref
Eisaguirre JH, Eisaguirre JM, Davis K, Carlson PM, Gaines SD, Caselle JE (2020) Trophic redundancy and predator size class structure drive differences in kelp forest ecosystem dynamics. Ecology 101:e02993 PubMed Crossref
Emmerson MC, Raffaelli D (2004) Predator-prey body size, interaction strength and the stability of a real food web. J Anim Ecol 73:399-409 Crossref
Estes JA, Duggins DO (1995) Sea otters and kelp forests in Alaska: generality and variation in a community ecological paradigm. Ecol Monogr 65:75-100 Crossref
Estes JA, Danner EM, Doak DF, Konar B and others (2004) Complex trophic interactions in kelp forest ecosystems. Bull Mar Sci 74:621-638
Estes JA, Terborgh J, Brashares JS, Power ME and others (2011) Trophic downgrading of Planet Earth. Science 333:301-306 Crossref
Eurich JG, Selden RL, Warner RR (2014) California spiny lobster preference for urchins from kelp forests: implications for urchin barren persistence. Mar Ecol Prog Ser 498:217-225 Crossref
Farina S, Oltra A, Boada J, Bartumeus F, Romero J, Alcoverro T (2018) Generation and maintenance of predation hotspots of a functionally important herbivore in a patchy habitat mosaic. Funct Ecol 32:556-565 Crossref
Fernandez LPH, Motta PJ (1997) Trophic consequences of differential performance: ontogeny of oral jaw-crushing performance in the sheepshead, Archosargus probatocephalus (Teleostei, Sparidae). J Zool 243:737-756 Crossref
Filbee-Dexter K, Scheibling R (2014) Sea urchin barrens as alternative stable states of collapsed kelp ecosystems. Mar Ecol Prog Ser 495:1-25 Crossref
FishBase (2025) Length-length relationship for Achoerodus viridis. www.fishbase.org Link
Flukes EB, Johnson CR, Ling SD (2012) Forming sea urchin barrens from the inside out: an alternative pattern of overgrazing. Mar Ecol Prog Ser 464:179-194
Freeman A (2006) Size-dependent trait-mediated indirect interactions among sea urchin herbivores. Behav Ecol 17:182-187 Crossref
Gabara SS, Konar BH, Edwards MS (2021) Biodiversity loss leads to reductions in community-wide trophic complexity. Ecosphere 12:e03361 Crossref
Galloway AWE, Gravem SA, Kobelt JN, Heady WN and others (2023) Sunflower sea star predation on urchins can facilitate kelp forest recovery. Proc R Soc B 290:20221897 PubMed Crossref
Giakoumi S, Pey A, Thiriet P, Francour P, Guidetti P (2019) Patterns of predation on native and invasive alien fish in Mediterranean protected and unprotected areas. Mar Environ Res 150:104792 PubMed Crossref
Gillanders BM (1995) Feeding ecology of the temperate marine fish Achoerodus viridis (Labridae): size, seasonal and site-specific differences. Mar Freshw Res 46:1009-1020 Crossref
Glasby TM, Gibson PT (2020) Decadal dynamics of subtidal barrens habitat. Mar Environ Res 154:104869 PubMed Crossref
Guidetti P (2006) Marine reserves reestablish lost predatory interactions and cause community changes in rocky reefs. Ecol Appl 16:963-976 PubMed Crossref
Guidetti P (2007) Predator diversity and density affect levels of predation upon strongly interactive species in temperate rocky reefs. Oecologia 154:513-520 PubMed Crossref PubMed Crossref
Guidetti P, Dulčić J (2007) Relationships among predatory fish, sea urchins and barrens in Mediterranean rocky reefs across a latitudinal gradient. Mar Environ Res 63:168-184
Gwinn DC, Allen MS, Johnston FD, Brown P, Todd CR, Arlinghaus R (2015) Rethinking length-based fisheries regulations: the value of protecting old and large fish with harvest slots. Fish Fish 16:259-281 Crossref
Hairston NG, Smith FE, Slobodkin LB (1960) Community structure, population control, and competition. Am Nat 94:421-425 Crossref
Hamilton SL, Caselle JE (2015) Exploitation and recovery of a sea urchin predator has implications for the resilience of southern California kelp forests. Proc R Soc B 282:20141817 Crossref
Hartig F (2017) DHARMa: residual diagnostics for hierarchical (multi-level / mixed) regression models. R package version 0.1.5. Link
Henry GW (1983) Biology and fisheries of yellowfin bream Acanthropagrus australis (Teleostei: Sparidae) in Tuggerah Lakes, New South Wales. Doctoral dissertation, UNSW Sydney
Hill NA, Blount C, Poore AGB, Worthington D, Steinberg PD (2003) Grazing effects of the sea urchin Centrostephanus rodgersii in two contrasting rocky reef habitats: effects of urchin density and its implications for the fishery. Mar Freshw Res 54:691 Crossref
Johnson CR, Banks SC, Barrett NS, Cazassus F and others (2011) Climate change cascades: shifts in oceanography, species’ ranges and subtidal marine community dynamics in eastern Tasmania. J Exp Mar Biol Ecol 400:17-32 Crossref Crossref
Johnson CR, Sanderson JC, Ling SD, Gardner C, Frusher SD, Redd KS, Pederson HG (2012) Identifying management options to minimise risk of development of sea-urchin barrens. In: Johnson C (ed) Echinoderms in a changing world: Proceedings of the 13th International Echinoderm Conference, January 5-9 2009, University of Tasmania, Hobart Tasmania, Australia. CRC Press, London, p 255
Kie JG (1999) Optimal foraging and risk of predation: effects on behavior and social structure in ungulates. J Mammal 80:1114-1129 Crossref
King C, Hoegh-Guldberg O, Byrne M (1994) Reproductive cycle of Centrostephanus rodgersii (Echinoidea), with recommendations for the establishment of sea urchin fishery in New South Wales. Mar Biol 120:95-106 Crossref
Klecka J, Boukal DS (2013) Foraging and vulnerability traits modify predator-prey body mass allometry: freshwater macroinvertebrates as a case study. J Anim Ecol 82:1031-1041 PubMed Crossref
Knott NA, Williams J, Harasti D, Malcolm HA and others (2021) A coherent, representative, and bioregional marine reserve network shows consistent change in rocky reef fish assemblages. Ecosphere 12:e03447 Crossref PubMed Crossref
Křivan V (1996) Optimal foraging and predator-prey dynamics. Theor Popul Biol 49:265-290
Kuiter RH (1993) Coastal fishes of south-eastern Australia. Crawford House Publishing, Goolwa, SA
Lennox RJ, Brownscombe JW, Darimont C, Horodysky A, Levi T, Raby GD, Cooke SJ (2022) The roles of humans and apex predators in sustaining ecosystem structure and function: contrast, complementarity and coexistence. People Nat 4:1071-1082 Crossref
Ling SD (2008) Range expansion of a habitat-modifying species leads to loss of taxonomic diversity: a new and impoverished reef state. Oecologia 156:883-894 PubMed Crossref
Ling S (2013) Pushing boundaries of range and resilience: a review of range-extension by a barrens-forming sea urchin. In: Fernández-Palacios JM, de Nascimento L, Hernández JC, Clemete S, González A, Díaz-González JP (ed) Climate change perspectives from the Atlantic: past, present and future. Servicio de Publicationes, Universidad de La Laguna, p 411-442
Ling SD, Johnson CR (2012) Marine reserves reduce risk of climate-driven phase shift by reinstating size- and habitat-specific trophic interactions. Ecol Appl 22:1232-1245 PubMed Crossref
Ling SD, Keane JP (2024) Climate-driven invasion and incipient warnings of kelp ecosystem collapse. Nat Commun 15:400 PubMed Crossref
Ling SD, Johnson CR, Frusher SD, Ridgway K (2009) Overfishing reduces resilience of kelp beds to climate-driven catastrophic phase shift. Proc Natl Acad Sci USA 106:22341-22345 PubMed Crossref
Ling SD, Johnson CR, Ridgway K, Hobday AJ, Haddon M (2009) Climate-driven range extension of a sea urchin: inferring future trends by analysis of recent population dynamics. Glob Change Biol 15:719-731 Crossref
Ling SD, Scheibling RE, Rassweiler A, Johnson CR and others (2015) Global regime shift dynamics of catastrophic sea urchin overgrazing. Philos Trans R Soc B 370:20130269 Crossref
Ling SD, Mahon I, Marzloff MP, Pizarro O, Johnson CR, Williams SB (2016) Stereo-imaging AUV detects trends in sea urchin abundance on deep overgrazed reefs. Limnol Oceanogr Methods 14:293-304 Crossref
Ling SD, Kriegisch N, Woolley B, Reeves SE (2019) Density-dependent feedbacks, hysteresis, and demography of overgrazing sea urchins. Ecology 100:e02577 PubMed Crossref
Litzow MA, Hunsicker ME (2016) Early warning signals, nonlinearity, and signs of hysteresis in real ecosystems. Ecosphere 7:e01614 Crossref
Luiz OJ, Crook DA, Kennard MJ, Olden JD and others (2019) Does a bigger mouth make you fatter? Linking intraspecific gape variability to body condition of a tropical predatory fish. Oecologia 191:579-585 PubMed Crossref
Mabuchi K, Miya M, Azuma Y, Nishida M (2007) Independent evolution of the specialized pharyngeal jaw apparatus in cichlid and labrid fishes. BMC Evol Biol 7:10 PubMed Crossref
MacDiarmid AB, Hickey B, Maller RA (1991) Daily movement patterns of the spiny lobster Jasus edwardsii (Hutton) on a shallow reef in northern New Zealand. J Exp Mar Biol Ecol 147:185-205 Crossref
Mann KH (1982) Ecology of coastal waters: a systems approach, Vol 8. University of California Press, Oakland, CA
Marinovich JK, Shears NT, Parsons DM (2025) Big fish, big mouths, big impacts: gape size of a fish predator determines size of urchins eaten. Mar Ecol Prog Ser 764:161-173 Crossref
Marzloff MP, Little LR, Johnson CR (2016) Building resilience against climate-driven shifts in a temperate reef system: staying away from context-dependent ecological thresholds. Ecosystems 19:1-15 Crossref
May JL, Maxwell JGH (1986) Trawl fish from temperate waters of Australia. CSIRO Division of Fisheries Research, Hobart
Mayfield S, De Beer E, Branch GM (2001) Prey preference and the consumption of sea urchins and juvenile abalone by captive rock lobsters (Jasus lalandii). Mar Freshw Res 52:773-780 Crossref
McClanahan TR, Muthiga NA (1989) Patterns of preedation [sic] on a sea urchin, Echinometra mathaei (de Blainville), on Kenyan coral reefs. J Exp Mar Biol Ecol 126:77-94 Crossref
McLaughlin RH, O’Gower AK (1971) Life history and underwater studies of a heterodont shark. Ecol Monogr 41:271-289 Crossref
Michel CJ, Henderson MJ, Loomis CM, Smith JM and others (2020) Fish predation on a landscape scale. Ecosphere 11:e03168 Crossref
Miller KI, Blain CO, Shears NT (2022) Sea urchin removal as a tool for macroalgal restoration: a review on removing ‘the spiny enemies’. Front Mar Sci 9:831001 Crossref
Morton JK, Platell ME, Gladstone W (2008) Differences in feeding ecology among three co-occurring species of wrasse (Teleostei: Labridae) on rocky reefs of temperate Australia. Mar Biol 154:577-592 Crossref
Motta PJ, Huber DR (2004) Prey capture behavior and feeding mechanics of elasmobranchs. In: Carrier JC, Musick JA, Heithaus MR (eds) Biology of sharks and their relatives. CRC Press, Boca Raton, FL, p 165-202
Myers RA, Worm B (2005) Extinction, survival or recovery of large predatory fishes. Philos Trans R Soc B 360:13-20 PubMed Crossref
Nichols KD, Segui L, Hovel KA (2015) Effects of predators on sea urchin density and habitat use in a southern California kelp forest. Mar Biol 162:1227-1237 Crossref
Nikanorov AM, Sukhorukov BL (2008) Ecological hysteresis. Dokl Earth Sci 423:1282-1285 Crossref
Norberg J (1999) Linking Nature’s services to ecosystems: some general ecological concepts. Ecol Econ 29:183-202 Crossref
NSW Department of Primary Industry (2023) Saltwater bag and size limits. Link
NSW Department of Primary Industry (2024) Eastern blue groper. Link
Paine RT (1980) Food webs: linkage, interaction strength and community infrastructure. J Anim Ecol 49:667-685 Crossref
Pederson HG, Johnson CR (2006) Predation of the sea urchin Heliocidaris erythrogramma by rock lobsters (Jasus edwardsii) in no-take marine reserves. J Exp Mar Biol Ecol 336:120-134 Crossref
Persson L, Andersson J, Wahlström E, Eklöv P (1996) Size-specific interactions in lake systems: predator gape limitation and prey growth rate and mortality. Ecology 77:900-911 Crossref
Pessarrodona A, Boada J, Pagès JF, Arthur R, Alcoverro T (2019) Consumptive and non-consumptive effects of predators vary with the ontogeny of their prey. Ecology 100:e02649 PubMed Crossref
Pine C, Erickson K, Gribben PE, Figueira WF (2022) Observation of juvenile eastern blue groper (Achoerodus viridis) on remnant oyster reefs in New South Wales, Australia. Ecology 103:e3824 PubMed Crossref
Powter DM, Gladstone W, Platell M (2010) The influence of sex and maturity on the diet, mouth morphology and dentition of the Port Jackson shark, Heterodontus portusjacksoni. Mar Freshw Res 61:74-85 Crossref
Preisser EL (2007) Trophic structure. In: Jorgensen SE (ed) Encyclopedia of ecology. Elsevier, Amsterdam, p 3608-3616
Pretorius JD, Lichtenstein JLL, Eliason EJ, Stier AC, Pruitt JN (2019) Predator-induced selection on urchin activity level depends on urchin body size. Ethology 125:716-723 PubMed Crossref
Przeslawski R, Chick R, Day J, Glasby T, Knott N (2023) Research summary—New South Wales barrens. NSW Department of Primary Industries, Nelson Bay
Pyke GH (1984) Optimal foraging theory: a critical review. Annu Rev Ecol Syst 15:523-575 Crossref
R Core Team (2023) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna
Ripple WJ, Estes JA, Schmitz OJ, Constant V and others (2016) What is a trophic cascade? Trends Ecol Evol 31:842-849 Crossref
Sala E (1997) Fish predators and scavengers of the sea urchin Paracentrotus lividus in protected areas of the north-west Mediterranean Sea. Mar Biol 129:531-539 Crossref
Sala E, Boudouresque CF, Harmelin-Vivien M (1998) Fishing, trophic cascades, and the structure of algal assemblages: evaluation of an old but untested paradigm. Oikos 82:425-439 Crossref
Sandin SA, Walsh SM, Jackson JBC (2010) Prey release, trophic cascades, and phase shifts in tropical nearshore ecosystems. In: Terborgh J, Estes JA (eds) Trophic cascades: predators, prey, and the changing dynamics of nature. Island Press, Washington, DC, p 71-90
Scharf FS, Juanes F, Rountree RA (2000) Predator size-prey size relationships of marine fish predators: interspecific variation and effects of ontogeny and body size on trophic-niche breadth. Mar Ecol Prog Ser 208:229-248 Crossref
Schiel DR (2013) The other 93%: trophic cascades, stressors and managing coastlines in non-marine protected areas. NZ J Mar Freshw Res 47:374-391
Schmitz O (2017) Predator and prey functional traits: understanding the adaptive machinery driving predator-prey interactions. F1000 Res 6:1767 PubMed Crossref
Segura AM, Franco-Trecu V, Franco-Fraguas P, Arim M (2015) Gape and energy limitation determine a humped relationship between trophic position and body size. Can J Fish Aquat Sci 72:198-205 Crossref
Selden RL, Gaines SD, Hamilton SL, Warner RR (2017) Protection of large predators in a marine reserve alters size-dependent prey mortality. Proc R Soc B 284:20161936 PubMed Crossref
Shears NT, Babcock RC (2002) Marine reserves demonstrate top-down control of community structure on temperate reefs. Oecologia 132:131-142 PubMed Crossref
Smith JE, Keane J, Oellermann M, Mundy C, Gardner C (2023) Lobster predation on barren-forming sea urchins is more prevalent in habitats where small urchins are common: a multi-method diet analysis. Mar Freshw Res 74:1493-1505 Crossref
Smith JE, Flukes E, Keane JP (2024) The risky nightlife of undersized sea urchins. Mar Freshw Res 75:MF23189 Crossref
Snellen CL, Hodum PJ, Fernández-Juricic E (2007) Assessing western gull predation on purple sea urchins in the rocky intertidal using optimal foraging theory. Can J Zool 85:221-231 Crossref
Sørdalen TK, Halvorsen KT, Olsen EM (2022) Protection from fishing improves body growth of an exploited species. Proc R Soc B 289:20221718 PubMed Crossref
Spyksma AJP, Taylor RB, Shears NT (2017) Predation cues rather than resource availability promote cryptic behaviour in a habitat-forming sea urchin. Oecologia 183:821-829 PubMed Crossref
Stachowicz JJ, Bruno JF, Duffy JE (2007) Understanding the effects of marine biodiversity on communities and ecosystems. Annu Rev Ecol Evol Syst 38:739-766 Crossref
Steneck RS (2020) Regular sea urchins as drivers of shallow benthic marine community structure. In: Lawrence JM (ed) Sea urchins: biology and ecology. Elsevier, London, p 255-279 Crossref
Stevenson CF, Demes KW, Salomon AK (2016) Accounting for size-specific predation improves our ability to predict the strength of a trophic cascade. Ecol Evol 6:1041-1053 PubMed Crossref
Stewart J (2023) NSW stock status summary 2021/22 - snapper (Chrysophrys auratus). Fisheries NSW, NSW Department of Primary Industries, Sydney Link
Stewart NL, Konar B (2012) Kelp forests versus urchin barrens: alternate stable states and their effect on sea otter prey quality in the Aleutian Islands. J Mar Sci 2012:492308
Stewart J, Rowling K, Hegarty AM, Nuttall A (2010) Size and age at sexual maturity of snapper Pagrus auratus in New South Wales 2008/09. Fish Res Rep Ser 27. Industry & Investment NSW, Cronulla Link
Stuart-Smith RD, Barrett NS, Crawford CM, Frusher SD, Stevenson DG, Edgar GJ (2008) Spatial patterns in impacts of fishing on temperate rocky reefs: Are fish abundance and mean size related to proximity to fisher access points? J Exp Mar Biol Ecol 365:116-125 Crossref
Taylor MD, McPhan L, van der Meulen DE, Gray CA, Payne NL (2013) Interactive drivers of activity in a free-ranging estuarine predator. PLOS ONE 8:e80962 PubMed Crossref
Teagle H, Hawkins SJ, Moore PJ, Smale DA (2017) The role of kelp species as biogenic habitat formers in coastal marine ecosystems. J Exp Mar Biol Ecol 492:81-98 Crossref
Therneau TM (1997) Extending the Cox model. In: Lin DY, Fleming TR (eds) Proceedings of the First Seattle Symposium in Biostatistics: survival analysis. Springer, New York, NY, p 51-84 Crossref
Therneau TM, Therneau MTM (2015) Package ’coxme’. R package version 2. Link
UNEP (United Nations Environment Programme) (2023) Into the blue: securing a sustainable future for kelp forests. www.unep.org/resources/report/blue-securing-sustainable-future-kelp-forests Link
van der Putten WH, de Ruiter PC, Martijn Bezemer T, Harvey JA, Wassen M, Wolters V (2004) Trophic interactions in a changing world. Basic Appl Ecol 5:487-494 Crossref
Vergés A, Tomas F, Ballesteros E (2012) Interactive effects of depth and marine protection on predation and herbivory patterns. Mar Ecol Prog Ser 450:55-65 Crossref
Wainwright PC, Smith WL, Price SA, Tang KL and others (2012) The evolution of pharyngognathy: a phylogenetic and functional appraisal of the pharyngeal jaw key innovation in labroid fishes and beyond. Syst Biol 61:1001-1027 PubMed Crossref
Weiss HM, Lozano-Álvarez E, Briones-Fourzán P (2008) Circadian shelter occupancy patterns and predator-prey interactions of juvenile Caribbean spiny lobsters in a reef lagoon. Mar Biol 153:953-963 Crossref
Wernberg T, Krumhansl K, Filbee-Dexter K, Pedersen MF (2019) Status and trends for the world’s kelp forests. In: Sheppard C (ed) World seas: an environmental evaluation, 2nd edn. Academic Press, London, p 57-78
Werner EE, Gilliam JF (1984) The ontogenetic niche and species interactions in size-structured populations. Annu Rev Ecol Syst 15:393-425 Crossref
Wood SN (2017) Generalized additive models: an introduction with R. Chapman and Hall/CRC, New York, NY Crossref
Woodward G, Ebenman B, Emmerson M, Montoya J, Olesen J, Valido A, Warren P (2005) Body size in ecological networks. Trends Ecol Evol 20:402-409 Crossref
Yatsuya K, Matsumoto Y (2023) Deterioration of an annual kelp Saccharina japonica forest and its effects on dominant herbivores, sea urchin Mesocentrotus nudus and abalone Haliotis discus hannai, in northeast Japan. Reg Stud Mar Sci 57:102739 Crossref
Young MAL, Foale S, Bellwood DR (2014) Impacts of recreational fishing in Australia. Environ Conserv 41:350-356 Crossref
Jessica Nguyen (Corresponding Author)
- Centre for Marine Science and Innovation, School of Biological, Earth & Environmental Sciences, UNSW Sydney, Kensington, NSW 2052, Australia
Scott Ling (Co-author)
- Institute for Marine & Antarctic Studies, University of Tasmania, Private Bag 129, Hobart, Tasmania 7001, Australia
John Keane (Co-author)
- Institute for Marine & Antarctic Studies, University of Tasmania, Private Bag 129, Hobart, TAS 7001, Australia
Adriana Vergés (Co-author)
- Centre for Marine Science and Innovation, School of Biological, Earth & Environmental Sciences, UNSW Sydney, Kensington, NSW 2052, Australia
Handling Editor:
John N. Griffin, Swansea, UK
Reviewers:
J. E. Caselle and 2 anonymous referees
Acknowledgements:
We thank Clayton Mead, Derrick Cruz, Sam Nolan, Tess Moriarty, Millie Brown, Jade McEwan, Olivier Pastore, Tahnee Chandler, Emmerson Sweeting, Nicole Sroba, Cata Musrri, Mitch Brennan and Suzy Evans for assisting with fieldwork. We thank Eve Slavich at Stats Central UNSW who advised on the statistical analysis of this study. We thank Andrew Niccum, Sergio Torres and Emily Bastow for help housing urchins in the Sydney Institute for Marine Science aquaria. Urchins were collected and deployed under permit P13/0007-2.0, and the experiment within an aquatic reserve was run under permit FP23/78-AR-MPL. This project was funded by ARC projects DP19102030 awarded to A.V. and FT200100949 to S.D.L. and by a WWF Innovate to Regenerate grant to A.V. Finally, we acknowledge that all fieldwork took place on the traditional lands of the Gadigal people of the Eora nation.
