Abstract
Purpose of the Review
The importance of habitat fragmentation in driving biodiversity loss has been recently debated. While the negative effects of habitat loss are well-documented, the effects of habitat fragmentation independent of habitat loss (e.g., habitat configuration) are more equivocal. Marine ecosystems have been underrepresented in past reviews, yet may differ fundamentally from terrestrial systems in their responses to habitat fragmentation because of the nature of energy/material flow, open population structure of most marine species, and narrow habitat extents. We conducted a systematic literature review and meta-analysis on the effects of habitat fragmentation in marine ecosystems.
Recent Findings
In our review of 180 studies from 28 articles, we found that habitat fragmentation effects were more often negative than positive, although the overall mean effect did not differ from zero. Interestingly, the mean effect was positive when the response was a measure of abundance, biodiversity, or population/ecosystem stability. Habitat fragmentation had overwhelmingly negative effects when it involved hydrological fragmentation. We found some support for the fragmentation threshold hypothesis via a weak negative relationship between habitat percent cover in the landscape and the habitat fragmentation effect.
Summary
Results of this review on the effects of habitat fragmentation in marine ecosystems are largely consistent with another recent review finding that habitat fragmentation (independent of habitat loss) does not have consistent, negative impacts on biodiversity, and in many cases may increase biodiversity. Future work should focus on factors driving this variability and employ multi-scale frameworks to test for congruence between patch- and landscape-scale studies.
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References
Papers of particular interest, published recently, have been highlighted as: • Of importance •• Of major importance
Wilcove DS, McLellan CH, Dobson AP. Habitat fragmentation in the temperate zone. Conserv Biol. 1986;6:237–56.
Saunders DA, Hobbs RJ, Margules CR. Biological consequences of ecosystem fragmentation: a review. Conserv Biol. 1991;5:18–32.
Reed BJ, Hovel KA. Seagrass habitat disturbance: how loss and fragmentation of eelgrass Zostera marina influences epifaunal abundance and diversity. Mar Ecol Prog Ser. 2006;326:133–43. https://doi.org/10.3354/meps326133.
Bostrom C, Jackson EL, Simenstad CA. Seagrass landscapes and their effects on associated fauna: a review. Estuar Coast Shelf Sci. 2006;68:383–403. https://doi.org/10.1016/j.ecss.2006.01.026.
Cushman SA. Effects of habitat loss and fragmentation on amphibians: a review and prospectus. Biol Conserv. 2006;128:231–40.
Fahrig L. Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst. 2003;34:487–515. https://doi.org/10.1146/annurev.ecolsys.34.011802.132419.
Brooks TM, Mittermeier RA, Mittermeier CG, Da Fonseca GA, Rylands AB, Konstant WR, et al. Habitat loss and extinction in the hotspots of biodiversity. Conserv Biol. 2002;16:909–23.
Gaston KJ, Blackburn TM, Goldewijk KK. Habitat conversion and global avian biodiversity loss. Proc R Soc Lond Ser B Biol Sci. 2003;270:1293–300.
Newbold T, Hudson LN, Hill SL, Contu S, Lysenko I, Senior RA, et al. Global effects of land use on local terrestrial biodiversity. Nature. 2015;520:45.
Lomolino MV. The species-area relationship: new challenges for an old pattern. Prog Phys Geogr. 2001;25:1–21.
•• Fahrig L. Ecological responses to habitat fragmentation per se. Annu Rev Ecol Evol Syst. 2017;48:1–23. https://doi.org/10.1146/annurev-ecolsys-110316-022612This review challenges current thinking by demonstrating that habitat fragmentation per se more often has positive effects on biodiversity than negative ones.
Fahrig L, Arroyo-Rodriguez V, Bennett JR, Boucher-Lalonde V, Cazetta E, Currie DJ, et al. Is habitat fragmentation bad for biodiversity? Biol Conserv. 2019;230:179–86. https://doi.org/10.1016/j.biocon.2018.12.026.
•• Fletcher RJ Jr, Didham RK, Banks-Leite C, Barlow J, Ewers RM, Rosindell J, et al. Is habitat fragmentation good for biodiversity? Biol Conserv. 2018;226:9–15 This article critiques findings of Fahrig 2017 and aims to provide evidence supporting negative effects of habitat fragmentation on biodiversity.
Ewers RM, Didham RK. Confounding factors in the detection of species responses to habitat fragmentation. Biol Rev. 2006;81:117–42. https://doi.org/10.1017/s1464793105006949.
Henle K, Davies KF, Kleyer M, Margules C, Settele J. Predictors of species sensitivity to fragmentation. Biodivers Conserv. 2004;13:207–51.
Zambrano J, G-L CM, Yeager L, Fortunel C, Cordeiro NB, Beckman NG. The effects of landscape fragmentation on plant functional diversity: what do we know so far? Oecologia. 2019; Accepted.
Umetsu F, Pardini R. Small mammals in a mosaic of forest remnants and anthropogenic habitats: evaluating matrix quality in an Atlantic forest landscape. Landsc Ecol. 2007;22:517–30.
Jules ES, Shahani P. A broader ecological context to habitat fragmentation: why matrix habitat is more important than we thought. J Veg Sci. 2003;14:459–64.
Fahrig L. Relative effects of habitat loss and fragmentation on population extinction. J Wildl Manag. 1997:603–10.
Fahrig L. When does fragmentation of breeding habitat affect population survival? Ecol Model. 1998;105:273–92.
Flather CH, Bevers M. Patchy reaction-diffusion and population abundance: the relative importance of habitat amount and arrangement. Am Nat. 2002;159:40–56.
Smith AC, Fahrig L, Francis CM. Landscape size affects the relative importance of habitat amount, habitat fragmentation, and matrix quality on forest birds. Ecography. 2011;34(1):103–13.
Bradford DF, Neale AC, Nash MS, Sada DW, Jaeger JR. Habitat patch occupancy by toads (Bufo punctatus) in a naturally fragmented desert landscape. Ecology. 2003;84(4):1012–23. https://doi.org/10.1890/0012-9658(2003)084[1012:hpobtb]2.0.co;2.
• Yeager LA, Keller DA, Burns TR, Pool AS, Fodrie FJ. Threshold effects of habitat fragmentation on fish diversity at landscapes scales. Ecology. 2016;97(8):2157–66. https://doi.org/10.1002/ecy.1449This study found support for the fragmentation threshold hypothesis in a marine ecosystem and seperates area and habitat fragmentation per se effects at the landscape scale.
Trzcinski MK, Fahrig L, Merriam G. Independent effects of forest cover and fragmentation on the distribution of forest breeding birds. Ecol Appl. 1999;9(2):586–93.
Parker M, Mac NR. Habitat loss and the habitat fragmentation threshold: an experimental evaluation of impacts on richness and total abundances using grassland invertebrates. Biol Conserv. 2002;105(2):217–29.
Hixon MA, Pacala SW, Sandin SA. Population regulation: historical context and contemporary challenges of open vs. closed systems. Ecology. 2002;83(6):1490–508.
Hedges LV, Olkin I Statistical methods for meta-analysis. Academic press; 2014.
Rosenberg MS, Rothstein HR, Gurevitch J. Effect sizes: conventional choices and calculations. In: Handbook of meta-analysis in ecology and evolution; 2013. p. 61–71.
Viechtbauer W. Conducting meta-analyses in R with the metafor package. J Stat Softw. 2010;36(3):1–48.
Team RC. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing; 2018.
Barton K. Multi-model inference. R package version 1.43.6. 2019.
Froese R, Pauly D. FishBase. World Wide Web electronic publication. www.fishbase.org, version (04/2019). 2019.
Palomares MLD, Pauly D. SeaLifeBase. World Wide Web electronic publication. www.sealifebase.org, version (08/2019). 2019.
Green BC, Smith DJ, Underwood GJC. Habitat connectivity and spatial complexity differentially affect mangrove and salt marsh fish assemblages. Mar Ecol Prog Ser. 2012;466:177–92. https://doi.org/10.3354/meps09791.
Acosta CA, Robertson DN. Diversity in coral reef fish communities: the effects of habitat patchiness revisited. Mar Ecol Prog Ser. 2002;227:87–96. https://doi.org/10.3354/meps227087.
Araujo MS, Langerhans RB, Giery ST, Layman CA. Ecosystem fragmentation drives increased diet variation in an endemic livebearing fish of the Bahamas. Ecol Evolution. 2014;4:3298–308. https://doi.org/10.1002/ece3.1140.
Arponen H, Bostrom C. Responses of mobile epifauna to small-scale seagrass patchiness: is fragmentation important? Hydrobiologia. 2012;680:1–10. https://doi.org/10.1007/s10750-011-0895-x.
Ault TR, Johnson CR. Spatial variation in fish species richness on coral reefs: habitat fragmentation and stochastic structuring processes. Oikos. 1998;82:354–64. https://doi.org/10.2307/3546976.
Bonin MC, Almany GR, Jones GP. Contrasting effects of habitat loss and fragmentation on coral-associated reef fishes. Ecology. 2011;92:1503–12. https://doi.org/10.1890/10-0627.1.
Cole VJ. Alteration of the configuration of bioengineers affects associated taxa. Mar Ecol Prog Ser. 2010;416:127–36. https://doi.org/10.3354/meps08772.
Crotty SM, Sharp SJ, Bersoza AC, Prince KD, Cronk K, Johnson EE, et al. Foundation species patch configuration mediates salt marsh biodiversity, stability and multifunctionality. Ecol Lett. 2018;21:1681–92. https://doi.org/10.1111/ele.13146.
Godbold JA, Bulling MT, Solan M. Habitat structure mediates biodiversity effects on ecosystem properties. Proc Royal Soc B Biol Sci. 2011;278:2510–8. https://doi.org/10.1098/rspb.2010.2414.
Godet L, Fournier J, Jaffre M, Desroy N. Influence of stability and fragmentation of a worm-reef on benthic macrofauna. Estuar Coast Shelf Sci. 2011;92(3):472–9. https://doi.org/10.1016/j.ecss.2011.02.003.
Goodsell PJ, Connell SD. Can habitat loss be treated independently of habitat configuration? Implications for rare and common taxa in fragmented landscapes. Mar Ecol Prog Ser. 2002;239:37–44. https://doi.org/10.3354/meps239037.
Grober-Dunsmore R, Frazer TK, Beets JP, Lindberg WJ, Zwick P, Funicelli NA. Influence of landscape structure, on reef fish assemblages. Landsc Ecol. 2008;23:37–53. https://doi.org/10.1007/s10980-007-9147-x.
Harwell HD, Posey MH, Alphin TD. Landscape aspects of oyster reefs: effects of fragmentation on habitat utilization. J Exp Mar Biol Ecol. 2011;409:30–41. https://doi.org/10.1016/j.jembe.2011.07.036.
Hattori A, Shibuno T. Total volume of 3D small patch reefs reflected in aerial photographs can predict total species richness of coral reef damselfish assemblages on a shallow back reef. Ecol Res. 2015;30:675–82. https://doi.org/10.1007/s11284-015-1268-0.
Hattori A, Shibuno T. The effect of patch reef size on fish species richness in a shallow coral reef shore zone where territorial herbivores are abundant. Ecol Res. 2010;25:457–68. https://doi.org/10.1007/s11284-009-0675-5.
Healey D, Hovel KA. Seagrass bed patchiness: effects on epifaunal communities in San Diego Bay, USA. J Exp Mar Biol Ecol. 2004;313:155–74. https://doi.org/10.1016/j.jembe.2004.08.002.
Hovel KA. Habitat fragmentation in marine landscapes: relative effects of habitat cover and configuration on juvenile crab survival in California and North Carolina seagrass beds. Biol Conserv. 2003;110:401–12. https://doi.org/10.1016/s0006-3207(02)00234-3.
Hovel KA, Wahle RA. Effects of habitat patchiness on American lobster movement across a gradient of predation risk and shelter competition. Ecology. 2010;91:1993–2002. https://doi.org/10.1890/09-0595.1.
Irlandi EA. Seagrass patch size and survivorship of an infaunal bivalve. Oikos. 1997;78:511–8. https://doi.org/10.2307/3545612.
Irlandi EA. Large-scale and small-scale effects of habitat structure on rates of predation: how percent coverage of seagrass affects rates of predation and siphon nipping on an infaunal bivalve. Oecologia. 1994;98:176–83. https://doi.org/10.1007/bf00341470.
Livernois MC, Grabowski JH, Poray AK, Gouhier TC, Hughes AR, O'Brien KF, et al. Effects of habitat fragmentation on Zostera marina seed distribution. Aquat Bot. 2017;142:1–9. https://doi.org/10.1016/j.aquabot.2017.05.006.
Ljungberg P, Hasper TB, Nilsson PA, Persson A. Effects of small-scale habitat fragmentation on predator-prey interactions in a temperate sea grass system. Mar Biol. 2013;160:667–75. https://doi.org/10.1007/s00227-012-2122-3.
Matias MG, Arenas F, Rubal M, Pinto IS. Macroalgal composition determines the structure of benthic assemblages colonizing fragmented habitats. PLoS One. 2015;10. https://doi.org/10.1371/journal.pone.0142289.
Morton DN, Shima JS. Habitat configuration and availability influences the settlement of temperate reef fishes (Tripterygiidae). J Exp Mar Biol Ecol. 2013;449:215–20. https://doi.org/10.1016/j.jembe.2013.09.017.
Ryan MR, Killen SS, Gregory RS, Snelgrove PVR. Predators and distance between habitat patches modify gap crossing behaviour of juvenile Atlantic cod (Gadus morhua, L. 1758). J Exp Mar Biol Ecol. 2012;422:81–7. https://doi.org/10.1016/j.jembe.2012.04.017.
Valentine-Rose L, Layman CA. Response of fish assemblage structure and function following restoration of two small Bahamian tidal creeks. Restor Ecol. 2011;19:205–15. https://doi.org/10.1111/j.1526-100X.2009.00553.x.
Valentine-Rose L, Layman CA, Arrington DA, Rypel AL. Habitat fragmentation decreases fish secondary production in Bahamian tidal creeks. Bull Mar Sci. 2007;80:863–77.
Leibold MA, Holyoak M, Mouquet N, Amarasekare P, Chase JM, Hoopes MF, et al. The metacommunity concept: a framework for multi-scale community ecology. Ecol Lett. 2004;7(7):601–13.
Cooper JK, Li J, Montagnes DJ. Intermediate fragmentation per se provides stable predator-prey metapopulation dynamics. Ecol Lett. 2012;15:856–63.
MacArthur RH. Patterns of species diversity. Biol Rev. 1965;40(4):510–33.
Pringle CM. Hydrologic connectivity and the management of biological reserves: a global perspective. Ecol Appl. 2001;11:981–98.
Haddad NM, Brudvig LA, Clobert J, Davies KF, Gonzalez A, Holt RD, et al. Habitat fragmentation and its lasting impact on Earth’s ecosystems. Sci Adv. 2015;1:e1500052.
•• Carroll JM, Keller DA, Furman BT, Stubler AD. Rough around the edges: lessons learned and future directions in marine edge effects studies. Curr Landsc Ecol Rep. 2019:Online first:1–12. This review finds that edge effects, a common patch-scale measure of habitat fragmentation, do not have consistent results of biodviersity in marine systems.
Acknowledgments
The authors thank Drs. Kevin Hovel and Lenore Fahrig for the invitation to contribute this review. We also thank Dr. Kevin Hovel for his thoughtful review which served to improve the manuscript.
Author Contribution Statement
Lauren Yeager conceptualized the study. All authors helped to review articles and extract data. Lauren Yeager analyzed the data and wrote the first draft of the manuscript on which all other authors provided feedback.
Funding
Lauren Yeager acknowledges funding support from the National Science Foundation award OCE # 1661683.
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Lauren Yeager, Jenelle Estrada, Kylie Holt, Spencer Keyser, and Tobi Oke declare that they have no conflict of interest.
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Table S1
Meta-data and effect sizes for all studies used in the review. (DOCX 49 kb)
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Yeager, L.A., Estrada, J., Holt, K. et al. Are Habitat Fragmentation Effects Stronger in Marine Systems? A Review and Meta-analysis. Curr Landscape Ecol Rep 5, 58–67 (2020). https://doi.org/10.1007/s40823-020-00053-w
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DOI: https://doi.org/10.1007/s40823-020-00053-w