Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

The community composition of Californian coastal sage scrub

Summary Coastal sage scrub is a community found extensively throughout cismontane California south of San Francisco, but has been surprisingly little studied. In the study area, which extends from Santa Barbara to the San Gorgonio Pass, two major floristic groupings can be found. In the basin bounded coastwards by a line drawn along the axis of the Santa Ana Mountains a large number of native and introduced annual herbs and a few shrubs (e.g.Encelia farinosa), rare or absent in the remainder of the study area, characterize one floristic group. In the coastal region the variety of shrub species increases, and the herbs are predominantly native and more restricted in number. Eleven groups defined by physiognomy, structure and species dominance, and arbitrarily called associations, are recognized. These associations can be grouped into four physiognomic-structural types which transgress the boundaries of the floristic groups. The results of this study and the limited previous literature suggest that Californian coastal sage scrub could be divided, mainly on floristic criteria, into Venturan, San Diegan and Riversidian sage.Plant nomenclature follows Munz & Keek (1968).We gratefully acknowledge the financial help provided by the Department of Earth Sciences, University of California, Riverside, and the aid in plant identification provided by Mr. Oscar Clarke, Museum Scientist, Department of Biology, University of California, Riverside.     

Xerie Mediterranean-type shrubland associations of Alta and Baja California and the community/continuum debate

A survey of the xeric shrublands of Pacific coastal North America from San Francisco to El Rosario (Mexico), including the inner Channel Islands, was conducted using 99 sample sites of 0.063 ha size. TWINSPAN classification and DECORANA ordination confirmed the existence of two plant formations, distinguishable physiognomically: coastal sage scrub and coastal succulent scrub. Within coastal sage scrub, four floristic associations were recognized: Diablan, Venturan, Riversidian and Diegan. Within coastal succulent scrub, two floristic associations were defined: Martirian and Vizcainan. These associations occur in distinct geographical regions following the coastline, with the Riversidian association occurring in the basin inland from Venturan and Diegan regions. Their locations are strongly correlated with differences in evapotranspirative stress regimes. The Channel Island sites show affinities to several of the mainland associations. The Venturan association can be further subdivided floristically into two subassociations, dominated by Salvia mellifera and S. leucophylla respectively. These subassociations which are coextensive geographically at a regional scale, typically do not intermingle at a local scale but often meet along sharp boundaries in the landscape. The dominant species segregate by moisture preference, S. mellifera preferring coarser-texture soils and more southerly aspects than S. leucophylla. Richness and equitability of these sites are depressed relative to other xeric shrubland sites, reflecting the fact that the two subassociations partition the Venturan flora into substantially non-overlapping subsets of species. This segregation of associates between the two Salvia dominance types strongly suggests biotic influence of the dominants on subordinate species, perhaps mediated by allelopathy. This biotic interaction, leading to relatively strong floristic subassociations segregating independently in the landscape, would provide an example of the holistic community structure referred to by Clements and his followers, embedded within a larger pattern of continuity in species distributions.Nomenclature follows Munz & Keck (1959), Munz (1974) and Wiggins (1980).I am grateful to the following for research assistance: S. Coon, E. Hobbs, S. Lavinger, J. F. O'Leary, K. R. Preston, B. Rich and A. Troeger. I also thank the numerous public and private land owners who permitted access to the study sites. This research is based upon work supported by the National Science Foundation under grant DEB 76-81712.     

Local lithological drivers of post-fire vegetation recovery and implications for fire-prone regions

Local ecosystem resilience to fire disturbance can be influenced by multiple factors, from topography and climate, to fire history and pre-fire structure of biotic communities. Here we investigated the factors affecting post-fire recovery of scrub vegetation in areas under Mediterranean climate affected by frequent fires. We hypothesized that, under comparable climatic and topographic conditions, geological factors (with bedrock type as a proxy) would be at least as important as fire history in explaining patterns of post-fire recovery. We surveyed scrub vegetation in a mountain study area in Portugal, using a stratified random sampling scheme, with fire frequency, time since last fire, and bedrock type (granite vs. schist) as stratifying layers. Based on vegetation and plant community data from 40 plots, we analyzed total species richness and composition, and the relative abundance of functional groups defined on the basis of general (non fire-specific) life-history traits. We found that, at a local scale, lithology can override fire history in determining post-fire recovery. Vegetation plots on granite exhibited a considerable development of tall scrubs and higher values of total species richness. They also hosted higher numbers of animal-dispersed woody species, of trees and tall scrubs, of woody deciduous species, and of forest, edge and tall scrub species. Differences in the post-fire development of scrub vegetation and in the functional profile of plant communities highlight the need to consider local geological diversity when establishing priorities for post-fire active restoration under scenarios of limited resources.     

Modelling the distribution of a threatened habitat: the California sage scrub

Aim Using predictive species distribution and ecological niche modelling our objectives are: (1) to identify important climatic drivers of distribution at regional scales of a locally complex and dynamic system – California sage scrub; (2) to map suitable sage scrub habitat in California; and (3) to distinguish between bioclimatic niches of floristic groups within sage scrub to assess the conservation significance of analysing such species groups. Location Coastal mediterranean‐type shrublands of southern and central California. Methods Using point localities from georeferenced herbarium records, we modelled the potential distribution and bioclimatic envelopes of 14 characteristic sage scrub species and three floristic groups (south‐coastal, coastal–interior disjunct and broadly distributed species) based upon current climate conditions. Maxent was used to map climatically suitable habitat, while principal components analysis followed by canonical discriminant analysis were used to distinguish between floristic groups and visualize species and group distributions in multivariate ecological space. Results Geographical distribution patterns of individual species were mirrored in the habitat suitability maps of floristic groups, notably the disjunct distribution of the coastal–interior species. Overlap in the distributions of floristic groups was evident in both geographical and multivariate niche space; however, discriminant analysis confirmed the separability of floristic groups based on bioclimatic variables. Higher performance of floristic group models compared with sage scrub as a whole suggests that groups have differing climate requirements for habitat suitability at regional scales and that breaking sage scrub into floristic groups improves the discrimination between climatically suitable and unsuitable habitat. Main conclusions The finding that presence‐only data and climatic variables can produce useful information on habitat suitability of California sage scrub species and floristic groups at a regional scale has important implications for ongoing efforts of habitat restoration for sage scrub. In addition, modelling at a group level provides important information about the differences in climatic niches within California sage scrub. Finally, the high performance of our floristic group models highlights the potential a community‐level modelling approach holds for investigating plant distribution patterns.     

Patterns of reproductive effort with time since last fire in Florida scrub plants

Abstract. In periodically burned ecosystems, fire frequency may be an important selective pressure for the evolution of plant reproductive allocation patterns. We evaluated this hypothesis for Florida (USA) scrub plants by developing three models of reproductive effort with time since last fire given assumptions concerning seed dormancy and seedling establishment. We then examined reproductive effort of five woody, resprouting shrubs at sites representing nine times since last fire (ranging from 0–64 yr). All species showed significant patterns with time since fire in percentage of stems reproductive and fruit production. Stems of all species needed to attain a minimum size before flowering. Four species had the greatest level of reproductive effort (fruit biomass/above-ground biomass) within 5 yr post-fire and best fit the Early Peak Model of reproductive effort (i.e. between-fire seedling recruitment or seed dormancy). A fifth species best fit the Broad Peak Model (i.e. immediate post-fire seedling establishment), peaking in reproductive effort at 7 yr post-fire. Both of these models are based on somewhat variable fire-return intervals, suggesting that the frequency of scrub fires may have been too unpredictable to select for reproductive allocation patterns precisely reflecting particular fire-return intervals. Early peaks in post-fire reproductive effort may be a bet-hedging strategy to allow for greater chances of seedling establishment and survival.     

白桦种群的稳定性与火干扰关系的研究

从种群生态学和干扰生态学的角度出发,在火史重建的基础上,采用种群世代划分的方法,研究了大兴安岭北部白桦群的稳定性与火干扰的关系,结果表明：白桦种群的耐火力与自我恢复力都较差,主要依靠很强的火后恢复力维持其稳定性,种群耐火力与长期火状况密切相关,表现为：低频类〈中频类〈高频类。高强类〈中强类〈低强类。火后恢复力与自我恢复力和最近一次火烧强度及距今时间密切相关,火后恢复力与自我恢复力都以低强类〉高强类     

The challenge of plant regeneration after fire in the Mediterranean Basin: scientific gaps in our knowledge on plant strategies and evolution of traits

Though observations on re-colonisation of post-fire sites in the Mediterranean Basin are plentiful, there still is an ongoing debate on the interrelation of fire regimes and species traits related to fire adaptation. Most of the studies found are restricted to particular species or claim to present community attributes. Therefore they often lack information for the evaluation of evolutionary evidence and historical contingency of the local fire regime and other abiotic conditions, which may act as selective pressure for plant regeneration strategies. Indeed, knowledge about the success of regeneration mechanisms and their interrelation with ecological factors is essential for the interpretation of the high spatio-temporal variability found in post-fire species performance. Such knowledge would be necessary to assess the potential of different regeneration mechanisms to cope with ongoing land-use and climate change—a crucial scientific challenge. A summary is given of the knowledge about the limits and potential of plant regeneration mechanisms after fire in the Mediterranean Basin, along with corresponding studies conducted in other parts of the world with similar climatic conditions in order to present the fullest possible picture. Moreover, the positive or negative impacts of particular parameters of a fire regime on different regeneration strategies (post-fire seeders, resprouters, and facultative resprouters) are explained and discussed in the light of published literature. To conclude, reference is made to scientific gaps that need to be filled in order to analyse species resistance and community resilience absorbing possible climate or land use changes.     

Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

兴安落叶松种群的稳定性与火干扰关系的研究

从种群生态学和干扰生态学的角度出发,采用火史重建的方法,研究了大兴安岭北部兴安落叶松种群的稳定性与火干扰的关系。结果表明：研究区内历史上火干扰频繁,平均间隔期为２８．９年,火烧强度较低。兴安落叶松依靠较强的耐火力、火力恢复力及自我恢复力具一定的稳定性。种群耐火力与长期火状况密切相关,表现为：低频类〈中频类〈高频类,高强类〈中强类〈低强类。火后恢复力与自我恢复力和最近一次火烧强度及距今时间密切相关：     

The Importance of Community Classification to Mitigation and Restoration of Coastal Sage Scrub

Community classification has emerged from ecology textbooks into the arena of environmental impact evaluation and mitigation. This development is especially apparent in southern California, where the fate of a community called coastal sage scrub is being decided. Regional plans for development, mitigation, preservation, and restoration are being formulated that will permanently affect the natural landscape. This paper demonstrates the potential for the name of a plant community to affect tradeoffs in planning processes that are intended to offset removal of habitat by development. The discussion focuses on two types of classification systems. One system is hierarchical, with established nomenclatorial rules that allow natural variation to refine the community definitions. Names of plant associations directly reflect the dominant species in the association, as in Artemisia californica—Eriogonum fasciculatum (California sagebrush—California buckwheat). The second system is nonhierarchical, in which observations of natural landscapes are fit into established definitions of community types, such as Diegan coastal sage scrub. Advantages and disadvantages of both types of systems are discussed and illustrated by two examples of problems in classifying coastal sage scrub. The discussion concludes with the point that the name of a community has the potential to significantly affect the extent to which equal tradeoffs between community types are actually achieved in mitigation and restoration efforts.     

Post-fire and post-quarry rehabilitation successions in Mediterranean-like ecosystems: Implications for ecological restoration

Resilience against sudden changes in the environment is a very desirable trait in plants used for ecosystem restoration. Mediterranean-like vegetation exhibits particularly strong fire resilience. There are two main functional groups of fire-prone species among Mediterranean-like vegetation: seeders and resprouters. Our aims were to describe how the theory of succession after fire relates to rehabilitation and to use this knowledge to improve the results of rehabilitation attempts in Mediterranean-like ecosystems. Eight post-fire (PF) sites, 14 post-rehabilitated (PR) quarry sites and two woodland sites were sampled. Detrended Canonical Correspondence Analysis (DCCA) showed that PF and PR successions were quite different. Both displayed an increasing abundance of resprouters over time, but seeder density increased throughout PR succession and decreased during PF succession. Nine species were common to both successions in all studied stages. The results showed that until 15-21 years of succession, the post-rehabilitation sites had not become as resilient to fires as sites populated by indigenous vegetation due to the lack of a seeder seed bank. However, after 21 years of PR succession, the exponentially increasing seeder population may allow for seed bank formation and thus eventually improve the fire resilience of the site.     

Fire history and patterns of Venturan subassociations of Californian coastal sage scrub

Californian coastal sage scrub has floristically distinct subassociations with sharp boundaries in the Santa Monica Mountains. This mesoscale biogeographic pattern has been variously attributed to the timing and pattern of fire and to differences in the moisture availability on sites. An examination of the actual fire history of sites reveals that recent fire events are unlikely to have caused the observed patterns. Sites with similar fire histories are not as similar in vegetation as sites with different fire histories but similar aspect. Single short fire intervals do not result in dissimilar communities; fires are unlikely to recur with the same boundaries as these two subassociations. A simulation of shrub response to fire interval and intensity indicates that long-term fire trends are also unlikely to have caused the distinction or the sharp boundaries of the two subassociations. Both subassociations include species with life history characteristics adapted to short and to long fire intervals. The subassociations exist without regard to fire history.     

Population structure and spatial pattern in the dioecious shrub Ceratiola ericoides

Abstract. The dioecious shrub Ceratiola ericoides (Florida rosemary) dominates xeric, infrequently burned Florida scrub vegetation, often to the near-exclusion of other woody species. We studied the spatial pattern, age, sex and size structure of four populations in Florida, USA: two coastal scrub populations subject to recurrent local disturbances due to sand movement, and two inland scrub populations in sites periodically burned by stand-replacing fires. The age structure of individual genets was estimated from node counts and used to describe the age structure of the populations. The sex ratio of males to females was not significantly different from 1:1, except within a female-biased coastal population subject to frequent sand movement. Node counts indicated that the mean age for reproductive individuals was 15 - 16 yr for the inland populations and 13 - 16 yr for the coastal populations. In all sites, there was no difference in mean age between males and females. Vegetative reproduction was uncommon except for the least-disturbed coastal population where 72 % of the reproductive individuals originated through layering. Individuals were generally randomly dispersed at the coastal sites, whereas significant aggregation of males and females occurred in the inland sites where the populations were initiated following fire. Seedling recruitment was continuous in the disturbed coastal scrub site, where 35% of the individuals were juveniles. Most juveniles were dispersed from 0.5 to 0.75 m around females. At one of the inland sites, where juveniles comprised 11% of the population, juveniles were clustered at 0.25 to 5.75 m around females. Coastal populations were all-aged, while inland populations were uneven-aged. Recruitment appears to follow periods of disturbance; infrequent fire in the inland populations and continuous sand movement on the coast are factors initiating recruitment.     

Foraging ecology of the California gnatcatcher deduced from fecal samples

The California gnatcatcher is a threatened species essentially restricted to coastal sage scrub habitat in southern California. Its distribution and population dynamics have been studied intensely, but little is known about its diet. We identified arthropod fragments in 33 fecal samples of the California gnatcatcher to gain insight into its foraging ecology and diet. Fecal samples were collected from adult males, adult females, fledglings, and nestlings. Leaf- and planthoppers (Homoptera) and spiders (Araneae) predominated numerically in samples. Spider prey was most diverse, with eight families represented. True bugs (Hemiptera) and wasps, bees, and ants (Hymenoptera) were only minor components of the gnatcatcher diet. Gnatcatcher adults selected prey to feed their young that was larger than expected given the distribution of arthropod size available in their environment, and chicks were provisioned with larger prey items and significantly more grasshoppers and crickets (Orthoptera) and spiders than adults consumed themselves. Both adults and young consumed more sessile than active prey. Further studies are needed to determine whether arthropods sampled in coastal sage scrub that are common in fecal samples are good indicators of California gnatcatcher habitat. Received: 30 December 1998 / Accepted: 28 April 1999     

Foliar stable carbon and nitrogen isotopes in woody Mediterranean species with different life form and post-fire regeneration

Wildfire is an important ecological disturbance factor in most Mediterranean ecosystems. In the Mediterranean Basin, most shrub species can regenerate after fire by resprouting or seeding. Here, we hypothesize that post-fire regenerative syndromes may potentially co-vary with traits directly related to functional properties involved in resource use. Thus, seeders with a shorter life span and smaller size would have lower water-use efficiency (WUE) than re-sprouting species and would take up nutrients such as nitrogen from more superficial parts of the soil. To test this hypothesis, we compared leaf ¹³C and ¹⁵N signatures from 29 co-existing species with different post-fire regeneration strategies. We also considered life form as an additional explanatory variable of the differences between post-fire regenerative groups. Our data support the hypothesis that seeder species (which mostly evolved in the Quaternary under a Mediterranean climate) have lower WUE and less stomatal control than non-seeders (many of which evolved under different climatic conditions in the Tertiary) and consequently greater consumption of water per unit biomass. This would be related to their smaller life forms, which tend to have lower WUE and shorter life and leaf lifespan. Differences in ¹⁵N also support the hypothesis that resprouters have deeper root systems than non-resprouters. The study supports the hypothesis of an overlap between plant functional traits and plant attributes describing post-disturbance resilience.     

Post-fire diversity patterns in two subassociations of Californian coastal sage scrub

Abstract. Coastal sage scrub exhibits floristically dissimilar subassociations on opposing aspects in the Santa Monica Mountains of southern California. Southward-facing (Venturan I) and northward-facing (Ventaran II) subassociations were sampled at two scales (1 m² and 625 m²) each spring for five years following a fire. Mature (10- and 28 yr-old) sites of these subassociation types were also sampled in the study area. The subshrub and herbaceous components that were most prevalent on northerly slopes following the first post-burn year contributed the most cover for all ages examined. Aspect-related differences in species richness were most apparent at the smaller sampling scale where they were equal the first post-burn year and varied inversely thereafter. At this scale, richness values were two times higher on older north-facing sites, largely due to the herbaceous component. South-facing sites displayed progressive decreases in equitability through time. This pattern reflects gradual reduction in overall species richness combined with progressive site domination by fewer species, as manifested by increasing dominance concentration values. By contrast equitability on north-facing sites fluctuated little through time, though dominance concentration tended to decrease. Higher richness, equitability, and overall cover with time after burning on north-facing sites is associated with the relatively mesic habitats upon which this subassociation develops. Post-burn patterns of species richness and stratal differentiation on south-facing slopes are consistent with those reported for other mediterranean-type shrublands, i.e. peak richness was attained the first two years following a fire with subsequent decrease to a constant level.     

Scale-dependent segregation of seeders and resprouters in cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis>) forests

Recent studies showed that disturbances and water availability determine the richness among plants with different post-fire strategies of Mediterranean-type ecosystems. The aim of this study was to determine whether or not the scale of analysis has an influence on the effects of these factors and, therefore, on the segregation of the dominant post-fire strategies, obligate seeders and obligate resprouters, and facultative species. We recorded all woody species and geographical features on 94 (75 m²) plots of cork oak woodlands in the southern Iberian Peninsula. For each regenerative type (resprouters, seeders and species with both traits—facultative species), we tested the relationship between the number of species and the predictors using a generalised linear mixed model. The fixed predictor considered at the large scale was altitude, and fixed predictors considered at the local scale were aspect (north/south) and disturbance (fire and clearing by heavy machinery; yes/no). The random predictor was the factor of site. When this factor did not have significant effect for some regenerative types, these relationships was tested using a generalised linear model. Resprouting species were most represented at lower altitudes and in undisturbed sites, while seeders were also at lower altitudes but mostly on south-facing slopes, especially south-facing disturbed sites. For facultative species, site is the most important variable. The proportion of seeders from the total species is not related to altitude, but it is related to disturbance and aspect. These results suggest that there is no segregation of the richness of seeders and resprouters at the large scale (altitudinal gradient). Differences appeared at the local scale (aspect and disturbance).     

Predicting Ecosystem Resilience to Fire from Tree Ring Analysis in Black Spruce Forests

Climate change has increased the occurrence, severity, and impact of disturbances on forested ecosystems worldwide, resulting in a need to identify factors that contribute to an ecosystem’s resilience or capacity to recover from disturbance. Forest resilience to disturbance may decline with climate change if mature trees are able to persist under stressful environmental conditions that do not permit successful recruitment and survival after a disturbance. In this study, we used the change in proportional representation of black spruce pre- to post-fire as a surrogate for resilience. We explored links between patterns of resilience and tree ring signals of drought stress across topographic moisture gradients within the boreal forest. We sampled 72 recently (2004) burned stands of black spruce in interior Alaska (USA); the relative dominance of black spruce after fire ranged from almost no change (high resilience) to a 90% decrease (low resilience). Variance partitioning analysis indicated that resilience was related to site environmental characteristics and climate–growth responses, with no unique contribution of pre-fire stand composition. The largest shifts in post-fire species composition occurred in sites that experienced the compounding effects of pre-fire drought stress and shallow post-fire organic layer thickness. These sites were generally located at warmer and drier landscape positions, suggesting they are less resilient to disturbance than sites in cool and moist locations. Climate–growth responses can provide an estimate of stand environmental stress to climate change and as such are a valuable tool for predicting landscape variations in forest ecosystem resilience.