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Chaparral Succession1 1 Chaparral Succession Abstract: Vegetation changes following fire usually result in a rapid return to the predisturbance species composition and structure. Regrowth takes Richard J. Vogl2 place by root-crown sprouting and seed reproduction. Succession is cyclic except in ecotones and favorable sites where it is linear. Fire serves as a growth stimulus to the persisting plants, contributes to seed germination, and is a plant waste remover and ecosystem renewer. Other factors that affect suc- cession are mountain building, landslides, erosion, soil development, climatic variability, and environ- mental stress. A complete discussion of chaparral succession and With few exceptions, fire is the initiator of a thorough review of the literature on the subject secondary succession in chaparral. Secondary succes- is presented by Hanes (1977). As a result, a review sion following fire is also without seral stages of the literature on chaparral succession will not (Hanes 1977). The temporary cover of annual fire- be attempted in this manuscript. Rather, certain followers (Armstrong 1977) and short-lived perennial highlights of plant succession will be again herbs (Keeley and Johnson 1977) and subshrubs does presented and re-emphasized. In addition, a concep- not modify the site to encourage a species invasion tual model of chaparral succession is advanced that or replacement. In about 2 to 5 years after a fire is intended to help explain the controlling factors almost all of these species cease growth (Vogl and and dynamics of succession. Schorr 1972), and the spaces they occupied are usually taken by the expanding canopies of the re- sprouting or regrowing chaparral shrubs (Ammirati Chaparral vs. Classical Succession 1967); the same species that dominated the preburn vegetation. The seeds of these herbaceous plants Classical plant succession (Vogl 1970, Horn 1974, apparently persist in place until their growth is 1976) is largely nonexistent in California chaparral, again initiated by fire (Sweeney 1956). The rapidity except possibly in transitional areas where chaparral and magnitude of the responses of these fire-follow- intergrades with other vegetation types. Missing in ers, along with the usual large sizes of the burned chaparral is the usual replacement series of various areas, preclude that the seeds of most of these plants or seral stages, with each species modifying species invade the burned sites from adjacent the site until it becomes unfavorable for that spe- unburned areas (Westman 1979). cies and thereby available for invasion by another (Horton and Kraebel 1955, Sweeney 1956, Patric and Successional replacement is apparently hampered Hanes 1964, Hanes and Jones 1967, Hanes 1971, Vogl by the strong vegetative habits of many chaparral and Schorr 1972, Biswell 1974, Keeley 1977, Keeley shrubs. Once a shrub occupies a site, it physically andLedler 1978). In most instances there is no dominates that site. Perennial plants with the species replacement that progresses until a stable ability to recover after top damage or removal of or climax vegetation is established (Hedrick 1951). stems have the potential to extend this dominance Even on virgin sites, the colonizing plants are also for long periods, perhaps for hundreds of years. components of the mature vegetation. Virgin sites Anything short of the destruction of the basal crown that are sometimes invaded by such species as annual of the root system results in the persistence of grasses, Selaginella mosses, coffee fern (Pellaea these chaparral shrubs. Removal of the aerial por- andromedaefolia), and perennial herbs do not appear tions of these perennials by fire is not just an to be particularly modified or enhanced to encourage adversity that these plants must overcome, but it successional replacement with subshrubs followed by also prevents-plant senescence by physiological woody plants. Growth that resembles the early stages rejuvenation and growth stimulation (Vogl 1980). of classical primary succession does not necessarily Factors that help to initially determine the chapar- lead to chaparral development, and primary succession ral species that will become established on a given is most often accomplished by the direct invasion of site are exposure of site, steepness of slope, soil a site by climax chaparral elements. New sites in moisture, elevation (Hanes 1977), and possibly the chaparral ecosystems are continuously produced by ash bed present after fire (Vogl and Schorr 1972). landslides which are triggered by rain or when slopes are undercut by runoff, or produced by instability Most chaparral shrubs also produce chemicals that as a result of tectonic activities. inhibit competing invaders which contribute to site dominance (McPherson and Muller 1969, Hanes 1977). The allelopathic chemicals produced by nonsprouting shrubs may extend chemical suppression of other 1 Presented at the Symposium on Dynamics and species after the nonsprouting shrubs have died, Management of Mediterranean-type Ecosystems, thus helping to ensure the re-establishment of the June 22-26, 1981, San Diego, California. same species on that site. It has been noted that after the short-lived bush poppy (Dendromecon rigida) 2 Professor of Biology, California State or certain species of Ceanothus die some 20 to 40 University, Los Angeles, Calif. 90032. years or more after a fire (Quick 1959, Hanes 1971, Gen. Tech. Rep. PSW-58. Berkeley, CA: Pacific Southwest Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture; 1982. 81 Keeley 1975), the spaces they occupied remain by long-term overgrazing and radical changes in fire open and unvegetated, or are only filled by the frequencies (Vogl 1977). Much of what is presently aerial portions of adjacent shrubs. classified as coastal sage scrub may have been either chaparral (Hanes 1977) or perennial grassland prior It has been generally assumed that among shrub to its degradation. Grazing impacts usually extend species that possess the ability to regrow vegeta- well beyond the areas being directly used by live- tively after fire, plants established from seeds stock as rodent and other wild herbivore populations seldom contribute substantial numbers of individuals are displaced and reconcentrated. Ranching opera- that become components of the mature vegetation tions also often reduce the predators of an area (Hanes 1977). Usually little seedling growth occurs thereby allowing prey populations to reach abnormal on sites dominated by shrubs capable of vegetative numbers that negatively impact the surrounding vege- reproduction (Keeley 1977). It has been hypothesized tation. that when seedlings do become established, they pro- vide a temporary plant cover that is preferentially Coastal sage scrub appears to be relatively browsed by herbivores until the resprouting shrubs stable, in that in many areas it is not undergoing again regain dominance (Vogl and Schorr 1972). It rapid vegetational changes and quickly reverting to appears that seed reproduction is only important as chaparral (Mooney 1977). It may be that these areas a replacement for shrubs that fail to recover vegeta- are in a state of shock stagnation, whereby the tively, or in places where the shrubs have been degraded vegetation persists or is self perpetuating eliminated by such things as a landslide or a very for an indefinite period with no evidence of species hot fire. replacement. Degraded California grasslands and many desert plant communities also do not show signs of An alternate explanation is that seed reproduction recovery indicating that the relative stability of is commonplace among resprouting perennials and con- these disclimax stages is commonplace among degraded tributes substantially to the mature vegetational ecosystems. Unless coastal sage scrub is not a prod- cover (Howe and Carothers 1980). If this assumption uct of degradation or the disturbances have not which contradicts most other studies is correct, then caused irreversible changes, such areas would be ex- it appears that chaparral shrubs are shorter lived pected to be eventually replaced by chaparral. and have higher replacement rates than suspected, since the typical high densities of chaparral shrubs The same floristic elements that characterize tend to otherwise physically saturate any given site coastal sage scrub also occur on small, localized with little or no space for successful seedling sites within undisturbed chaparral. They usually establishment (Hanes 1977). Further research is respond to minor soil disturbances caused by ground needed on this subject, particularly since shrub squirrels, pocket gophers, rock slides, erosion, and seedlings might often be misidentified as resprouts the like. Under such disturbance conditions, these and vice versa (Howe and Carothers 1980). species act like pioneer or weedy species responding to the open and unstable conditions. These species Chaparral succession is also simplified by the are replaced by the more stable chaparral species absence of an understory vegetation (Hanes 1971, when the disturbances are eliminated. This type of 1977). The mature chaparral usually consists of a successional replacement is similar to that found in single, although sometimes irregular, layer or stra- perennial grasslands where short-lived opportunistic tum of dense shrub growth with canopies that are pioneers coexist with long-lived perennials. A continuous and contiguous with each other. The natural grassland is constantly being turned over, a general absence
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