Abstract
Context
This study provides a unified, holistic framework for predicting the dynamics of shrub-grass conversion throughout Mediterranean-climate shrublands. This work focuses specifically on the California chaparral, which until recently has been considered resistant to invasion by exotic grasses, but in recent years appears to have undergone substantial type conversion.
Objectives
To synthesize current understanding of the feedbacks and anthropogenic impacts that both enhance and reduce the susceptibility of southern California chaparral to invasion and its corresponding ability to recolonize invaded areas.
Methods
We review the existing literature pertaining to the factors that enhance or reduce the susceptibility of chaparral to invasion, and organize these factors and their interactions into a single unified framework of environmental drivers, ecological interactions, and historical legacies associated with the distribution and rate of such invasion.
Results
A myriad of processes interact to mediate the invasion of exotic grasses into intact chaparral. In addition, we demonstrate that feedbacks exist within both chaparral shrublands and exotic grasslands that modify the landscape in ways that can enhance their own survival, act as barriers to conversion into alternate cover types, and in some cases weaken the resistance of adjacent vegetation to invasion. We posit a methodological framework from which the many climatic, anthropogenic, edaphic, and biotic feedbacks that determine the mosaic of invasion can be modeled.
Conclusions
This study demonstrates that substantial conversion of chaparral into deciduous grasslands has recently occurred in southern California and presents a unified framework for forecasting the dynamics of shrub-grass conversion throughout Mediterranean-climate shrublands.
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Data availability
Example code used to produce estimates of herbaceous cover in this study are available through: https://doi.org/10.6084/m9.figshare.6203366.
References
Agee JK, Skinner CN (2005) Basic principles of forest fuel reduction treatments. For Ecol Manage 211:83–96
Allen EB (1999) Restoring habitats to prevent exotics. In: Proceedings of the California exotic pest plant council symposium, pp 41–44
Allen EB, Padgett PE, Bytnerowicz A, Minnich R (1998) Nitrogen deposition effects on coastal sage vegetation of southern California. In: Proceedings of the international symposium on air pollution and climate change effects on forest ecosystems, Pacific Southwest Research Station, Riverside, California, USA. US Department of Agriculture Forest Service, pp 131–140
Allen EB, Williams K, Beyers JL, Phillips M, Ma S, D’Antonio CM (2018) Chaparral Restoration. In: Underwood EC, Safford HD, Molinari NA, Keeley JE (eds) Valuing chaparral: ecological, socio-economic, and management perspectives. Springer, Berlin
Bartholomew B (1970) Bare zone between California shrub and grassland communities: the role of animals. Science 170:1210–1212
Bentley JR (1967) Conversion of Chaparral areas to grassland: techniques used in California vol Agriculture Handbook No. 328. Forest Service, USDA, Berkeley, CA
Borchert MI, Odion DC (1995) Fire intensity and vegetation recovery in chaparral: a review. In: Keeley JE, Scott T (eds) Brushfires in California wildlands: ecology and resource management. International Association of Wildland Fire, Fairfield
Bradshaw LS, Deeming JE, Burgan RE, Cohen JD (1983) The 1978 national fire-danger rating systemL technical documentation. United Stated Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station, Ogden
Brennan TJ, Keeley JE (2015) Effect of mastication and other mechanical treatments on fuel structure in chaparral. Int J Wildland Fire 24:949–963
Brooks ML (2002) Peak fire temperatures and effects on annual plants in the Mojave desert. Ecol Appl 12:1088–1102
Brooks ML, D'Antonio CM, Richardson DM, Grace JB, Keeley JE, DiTomaso JM, Hobbs RJ, Pellant M, Pyke D (2004) Effects of invasive alien plants on fire regimes. Bioscience 54:677–688
Bytnerowicz A, Johnson RF, Zhang L, Jenerette GD, Fenn ME, Schilling SL, Gonzalez-Fernandez I (2015) An empirical inferential method of estimating nitrogen deposition to Mediterranean-type ecosystems: the San Bernardino Mountains case study. Environ Pollut 203:69–88
Christensen NL (1985) Shrubland fire regime and their evolutionary consequences. In: Pickett STA (ed) The ecology of natural disturbance and patch dynamics. Academic Press Inc, Orlando, pp 85–99
Christensen NL, Muller CH (1975a) Effects of fire on factors controlling plant growth in adenostoma chaparral. Ecol Monogr 45:29–55
Christensen NL, Muller CH (1975b) Relative importance of factors controlling germination and seedling survival in adenostoma chaparral. Am Midl Nat 93:71–78
Conard SG, Weise DR (1998) Management of fire regime, fuels and fire effects in southern California chaparral: lessons from the past and thoughts for the future vol 20. Fire in ecosystem management: shiftng the paradigm from suppression to prescription. Tall Timbers Fire Ecology Conference Proceedings. Tall Timber Research Station, Tallahassee, Florida, USA
Corbett ES, Crouse RP (1968) Rainfall interception by annual grass and chaparral… losses compared (U.S. Forest Serv. Res. Paper PSW-48). USDA Forest Service, Pacific Southwest Forest and Range Experiment Station, Berkeley, California
Cornwell WK, Cornelissen JHC, Amatangelo K, Dorrepaal E, Eviner VT, Godoy O, Hobbie SE, Hoorens B, Kurokawa H, Pérez-Harguindeguy N, Quested HM, Santiago LS, Wardle DA, Wright IJ, Aerts R, Allison SD, Van Bodegom P, Brovkin V, Chatain A, Callaghan TV, Díaz S, Garnier E, Gurvich DE, Kazakou E, Klein JA, Read J, Reich PB, Soudzilovskaia NA, Vaieretti MV, Westoby M (2008) Plant species traits are the predominant control on litter decomposition rates within biomes worldwide. Ecol Lett 11:1065–1071
Costello DA, Lunt ID, Williams JE (2000) Effects of invasion by the indigenous shrub Acacia sophorae on plant composition of coastal grasslands in south-eastern Australia. Biol Conserv 96:113–121
Cox RD, Allen EB (2008) Stability of exotic annual grasses following restoration efforts I nsouthern California coastan sage scrub. J Appl Ecol 45:495–504
D’Antonio CM, Vitousek PM (1992) Biological invasions by exotic grasses, the grass/fire cycle, and global change. Annu Rev Ecol Syst 23:63–87
Davies KW, Nafus AM, Madsen MD (2013) Medusahead Invasion along unimproved roads, animal trails, and random transects. West N Am Nat 73:54–59
Davis EA (1984) Conversion of Arizona chaparral to grass increases water yield and nitrate loss. Water Resour Res 20:1643–1649
Davis EA (1987) Chaparral conversion and streamflow: nitrate increase is balanced mainly by a decrease in bicarbonate. Water Resour Res 23:215–224
Davis EA, Debano LF (1986) Nitrate increases in soil water following conversion of chaparral to grass. Biogeochemistry 2:53–65
Davis SD, Mooney HA (1985) Comparative water relations of adjacent California shrub and grassland communities. Oecologia 66:522–529
DiTomaso JM, Brooks ML, Allen EB, Minnich R, Rice PM, Kyser GB (2006) Control of invasive weeds with prescribed burning. Weed Technol 20:535–548
Dunn PH, Barro SC, Wells WG, Poth MA, Wohlgemuth PM, Colver CG (1988) The San Dimas experimental forest: 50 years of research. United States Department of Agriculture, Pacific Southwest Forest and Range Experiment Station, Albany
Eliason SA, Allen EB (1997) Exotic grass competition in suppressing native shrubland re-establishment. Restor Ecol 5:245–255
Fenn ME, Jovan S, Yuan F, Geiser LH, Meixner T, Gimeno BS (2008) Empirical and simulated critical loads for nitrogen deposition in California mixed conifer forests. Environ Pollut 155:492–511
Fenn ME, Allen EB, Weiss SB, Jovan S, Geiser LH, Tonnesen GS, Johnson RF, Rao LE, Gimeno BS, Yuan F, Meixner T, Bytnerowicz A (2010) Nitrogen critical loads and management alternatives for N-impacted ecosystems in California. J Environ Manage 91:2404–2423
Figueroa JA, Castro SA, Marquet PA, Jaksic FM (2004) Exotic plant invasions to the mediterranean region of Chile: causes, history and impacts. Rev Chil Hist Nat 77:465–483
Frazer JM, Davis SD (1988) Differential survival of chaparral seedlings during the first summer drought after wildfire. Oecologia 76:215–221
Gómez-González S, Torres-Díaz C, Valencia G, Torres-Morales P, Cavieres LA, Pausas JG (2011) Anthropogenic fires increase alien and native annual species in the Chilean coastal matorral. Divers Distrib 17:58–67.
Goodridge BM, Hanan EJ, Aguilera R, Wetherley EB, Chen Y-J, D’Antonio CM, Melack JM (2018) Retention of nitrogen following wildfire in a Chaparral. Ecosystems. https://doi.org/10.1007/s10021-018-0243-3
Gray JT, Schlesinger WH (1983) Nutrient use by evergreen and deciduous shrubs in Southern California: II. Experimental investigations of the relationship between growth, nitrogen uptake and nitrogen availability. J Ecol 71:43–56
Green LR (1977) Fuelbreaks and other fuel modification for wildland fire control. In: Agriculture handbook vol 499. USDA Forest Service, Washington DC
Grigulis K, Lavorel S, Davies ID, Dossantos A, Lloret F, Vilàs M (2005) Landscape-scale positive feedbacks between fire and expansion of the large tussock grass, Ampelodesmos mauritanica in Catalan shrublands. Glob Change Biol 11:1042–1053
Gritti ES, Smith B, Sykes MT (2006) Vulnerability of Mediterranean Basin ecosystems to climate change and invasion by exotic plant species. J Biogeogr 33:145–157
Haidinger TL, Keeley JE (1993) Role of high fire frequency in destruction of mixed chaparral. Madroño 40:141–147
Hedrick DW (1951) Studies on the succession and manipulation of chamise brushlands in California. Texas A&M University, Texas
Hibbert AR (1971) Increases in streamflow after converting chaparral to grass. Water Resour Res 7:71–80
Holmgren M (2002) Exotic herbivores as drivers of plant invasion and switch to ecosystem alternative states. Biol Invasions 4:25–33
Homyak PM (2012) Nitrogen and phosphorus biogeochemistry of watersheds along the western slope of the Sierra Nevada. UC Riverside
Huenneke LF, Hamburg SP, Koide R, Mooney HA, Vitousek PM (1990) Effects of soil resources on plant invasion and community structure in Californian serpentine grasslands. Ecology 71:478–491
Johnson WH, Fitzhugh EL (1990) Grazing helps maintain brush growth on cleared land California. Agriculture 44:31–35
Keeley JE (1992) Recruitment of seedlings and vegetative sprouts in unburned Chaparral. Ecology 73:1194–1208
Keeley JE (2000) Chaparral. In: Barbour M, Billings W (eds) North American terrestrial vegetation. Cambridge University Press, New York
Keeley JE (2001) Fire and invasive species in Mediterranean-climate ecosystems of California. In: Galley KEM, Wilson TP (eds) Proceedings of the invasive species workshop: the role of fire in the control and spread of invasive species. Timbers Research Station, Tallahassee, pp 81–94
Keeley JE (2004) Invasive plants and fire management in California Mediterranean-climate ecosystems. In: Proceedings 10th MEDECOS Conference
Keeley JE, Brennan TJ (2012) Fire-driven alien invasion in a fire-adapted ecosystem. Oecologia 169:1043–1052
Keeley JE, Keeley SC (1987) Role of fire in the germnation of chaparral herbs and suffrutescents. Madroño 34:240–249
Keeley JE, Syphard AD (2018) South Coast Bioregion. In: Wagtendonk JW, Sugihara NG, Stephens SL, Thode AE, Shaffer KE, Fites-Kaufman JA (eds) Fire in California’s ecosystems, 2nd edn. University of California Press, Berkeley, California
Keeley JE, Zedler PH (1978) Reproduction of chaparral shrubs after fire: a comparison of sprouting and seeding strategies. Am Midl Nat 99:142–161
Keeley JE, Morton BA, Pedrosa A, Trotter P (1985) Role of allelopathy, heat and charred wood in the germination of chaparral herbs and suffrutescents. J Ecol 73:445–458.
Keeley JE, Lubin D, Fotheringham CJ (2003) Fire and grazing impacts on plant diversity and alien plant invasions in the southern sierra nevada. Ecol Appl 13:1355–1374
Keeley JE, Fotheringham CJ, Baer-Keeley M (2005a) Determinants of postfire recovery and succession in mediterranean-climate shrublands of California. Ecol Appl 15:1515–1534
Keeley JE, Fotheringham CJ, Baer-Keeley M (2005b) Factors affecting plant diversity during post-fire recovery and succession of mediterranean-climate shrublands in California, USA. Divers Distrib 11:525–537
Kelly M, K-i Ueda, Allem-Diez B (2008) Considerations for ecological reconstruction of historic vegetation: analysis of the spatial uncertainties in the California vegetation type map dataset. Plant Ecol 194:37–49
Knops JMH, Griffin JR, Royalty AC (1995) Introduced and native plants of the Hastings Reservation, central coastal California: a comparison. Biol Conserv 71:115–123
Lambrinos JG (2002) The variable success of Cortaderia species in a complex landscape. Ecology 83:518–529
Lambrinos JG (2006) Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass. Oecologia 147:327–334
Lett M, Knapp AK (2003) Consequences of shrubexpansion in mesic grassland: resource alterations and graminoid responses. J Veg Sci 14:487–496
Lippitt C, Stow D, O’Leary J, Franklin J (2012) Influence of short-interval fire occurrence on post-fire recovery of fire-prone shrublands in California, USA. Int J Wildland Fire 22(2):184–193
Longstreth DT, Patten DT (1975) Conversion of chaparral to grass in central Arizona: effects on selected ions in watershed runoff. Am Midl Nat 93:25–34
McMaster GS, Jow WM, Kummerow J (1982) Response of Adenostema fasciculatum and Ceanothus greggii chaparral to nutrient additions. J Ecol 70:745–756
McPherson JK, Muller CH (1969) Allelopathic effects of Adenostoma fasciculatum, “Chamise”, in the California chaparral. Ecol Monogr 39:177–198
Merriam KE, Keeley JE, Beyers JL (2006) Fuel breaks affect nonnative species abundance in californian plant communities. Ecol Appl 16:515–527
Minnich RA, Bahr CJ (1995) Wildland fire and chaparral succession along the Californis Baja-California boundary. Int J Wildland Fire 5:13–24
Mooney HA, Rundel PW (1979) Nutrient relations of the evergreen shrub, Adenostoma fasciculatum, in the California Chaparral. Bot Gaz 140:109–113
Moreno JM, Oechel WC (1991) Fire intensity effects on germination of shrubs and herbs in southern California. Ecology 72:1993–2004
Muller RN (1982) Vegetation patterns in the mixed mesophytic forest of eastern Kentucky. Ecology 63:1901–1917
Muñoz MR, Fuentes ER (1989) Does fire induce shrub germination in the Chilean Matorral? Oikos 56:177–181
Murphy AH, Leonard OA (1974) Chaparral shrub control as influenced by grazing, herbicides and fire California. Agriculture 28:10–13
Neary DG, Klopatek CC, DeBano LF, Ffolliott PF (1999) Fire effects on belowground sustainability: a review and synthesis. For Ecol Manage 122:51–71
Odion DC, Davis FW (2000) Fire, soil heating, and the formation of vegetation patterns in chaparral. Ecol Monogr 70:149–169
Oechel WC (1988) Seedling establishment and water relations after fire in a Mediterranean ecosystem. In: Allen EB (ed) The reconstruction of disturbed arid lands. Westview Press, Boulder, pp 34–45
Orme AR, Bailey RG (1970) The effect of vegetation conversion and flood discharge on stream channel geometry: a case of southern California watersheds. Proc Am Assoc Geogr 2:101–106
Orrock JL, Witter MS, Reichman OJ (2008) Aparent competition with an exotic plant reduces native plant establishment. Ecology 89:1168–1174
Park I, Hooper J, Flegal JM, Jenerette GD (2018) Impacts of climate, disturbance, and topography on distribution of herbaceous cover in Southern California chaparral: insights from a remote sensing method. Divers Distrib 24:497–508
Parker VT (1987) Effects of wet-season management burns on chaparral vegetation: implications for rare species. In: Elias TE (ed) Proceedings of a conference on the conservation and management of rare and endangered plants, Sacramento. California Native Plant Society, California, pp 233–237
Rowe PB (1963) Streamflow increases after removing woodland-riparian vegetation from a southern california watershed. J For 61:365–370
Rundel PW (1983) Impact of fire on nutrient cycle. In: Kruger FJ, Mitchell DT, Jarvis JUM (eds) Mediterranean type ecosystems, vol 192–207. Springer, Berlin
Schultz AM, Launchbaugh JL, Biswell HH (1955) Relationship between grass density and brush seedling survival. Ecology 36:226–238
Segura AM, Holmgren M, Anabalón JJ, Fuentes ER (1998) The significance of fire intensity in creating local patchiness in the Chilean matorral. Plant Ecol 139:259–264
Stylinski CD, Allen EB (1999) Lack of native species recovery following severe exotic disturbance in southern Californian shrublands. J Appl Ecol 36:544–554
Syphard AD, Radeloff VC, Keeley JE, Hawbaker TJ, Clayton MK, Stewart SI, Hammer RB (2007) Human influence on California fire regimes. Ecol Appl 17:1388–1402
Syphard AD, Brennan TJ, Keeley JE (2019) Drivers of chaparral type conversion to herbaceous vegetation in coastal Southern California. Divers Distrib 25:90–101
Tyler CM (1995) Factors contributing to postfire seedling establishment in chaparral: direct and indirect effects of fire. J Ecol 83:1009–1020
Tyler CM (1996) Relative importance of factors contributing to postfire seedling establishment in maritime Chaparral. Ecology 77:2182–2195
Valliere JM, Irvine IC, Santiago L, Allen EB (2017) High N, dry: experimental nitrogen deposition exacerbates native shrub loss and nonnative plant invasion during extreme drought. Glob Change Biol 23:4333–4345
Van de Water K, Safford HD (2011) A summary of fire frequency estimates for California vegetation before Euro-American settlement fire. Ecology 7:26–58
van Wilgen BW, Richardson DM (1985) The effects of alien shrub invasions on vegetation structure and fire behaviour in South African Fynbos Shrublands: a simulation study. J Appl Ecol 22:955–966
Vilà M, Lloret F, Ogheri E, Terradas J (2001) Positive fire-grass feedback in Mediterranean Basin woodlands. For Ecol Manage 147:3–14
Vourlitis GL (2017) Chronic N enrichment and drought alter plant cover and community composition in a Mediterranean-type semi-arid shrubland. Oecologia 184:267–277
Vourlitis GL, Pasquini SC, Mustard JF (2009) Effects of dry-season N input on the productivity and N storage of Mediterranean-type shrublands. Ecosystems 12:473–488
Weiss SB (1999) Cars, cows, and checkerspot butterflies: nitrogen deposition and management of nutrient-poor grasslands for a threatened species. Conserv Biol 13(6):1476–1486
Williamson TN, Graham RC, Shouse PJ (2004a) Effects of a chaparral-to-grass conversion on soil physical and hydrologic properties after four decades. Geoderma 123:99–114
Williamson TN, Newman BD, Graham RC, Shouse PJ (2004b) Regolith water in zero-order chaparral and perennial grass watersheds four decades after vegetation conversion. Vasose Zone J 3:1007–1016
Wolkovich EM, Bolger DT, Cottingham KL (2009) Invasive grass litter facilitates native shrubs through abiotic effects. J Veg Sci 20:1121–1132
Wolkovich EM, Lipson DA, Virginia RA, Cottingham KL, Bolger DT (2010) Grass invasion causes rapid increases in ecosystem carbon storage in a semiarid shrubland. Glob Change Biol 16:1351–1365
Yoshida LC, Allen EB (2004) 15 N uptake by mycorrhizal native and invasive plants from a N-eutrophied shrubland: a greenhouse experiment. Biol Fertil Soils 39:243–248
Zammit CA, Zedler PH (1988) The influence of dominant shrubs, fire, and time since fire on soil seed banks in mixed chaparral. Vegetatio 75:175–187
Zedler PH, Gautier CR, McMaster GS (1983) Vegetation change in response to extreme events: the effect of a short interval between fires in California chaparral and coaster scrub. Ecology 64:809–818
Zink TA, Allen MF, Heinde-Tenhunen B, Allen EB (1995) The effect of a disturbance corridor on an ecological reserve. Restor Ecol 3:304–310
Acknowledgements
This research was funded through a cooperative agreement between UC-Riverside and the US Forest service (Grant No. 11-CS-11050100-031) and the National Science Foundation (DEB—1656062).
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Park, I.W., Jenerette, G.D. Causes and feedbacks to widespread grass invasion into chaparral shrub dominated landscapes. Landscape Ecol 34, 459–471 (2019). https://doi.org/10.1007/s10980-019-00800-3
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DOI: https://doi.org/10.1007/s10980-019-00800-3