Maritime climate influence on chaparral composition and diversity in the coast range of central California

We investigated the hypothesis that maritime climatic factors associated with summer fog and low cloud stratus (summer marine layer) help explain the compositional diversity of chaparral in the coast range of central California. We randomly sampled chaparral species composition in 0.1‐hectare plots along a coast‐to‐interior gradient. For each plot, climatic variables were estimated and soil samples were analyzed. We used Cluster Analysis and Principle Components Analysis to objectively categorize plots into climate zone groups. Climate variables, vegetation composition and various diversity measures were compared across climate zone groups using ANOVA and nonmetric multidimensional scaling. Differences in climatic variables that relate to summer moisture availability and winter freeze events explained the majority of variance in measured conditions and coincided with three chaparral assemblages: maritime (lowland coast where the summer marine layer was strongest), transition (upland coast with mild summer marine layer influence and greater winter precipitation), and interior sites that generally lacked late summer water availability from either source. Species turnover (β‐diversity) was higher among maritime and transition sites than interior sites. Coastal chaparral differs from interior chaparral in having a higher obligate seeder to facultative seeder (resprouter) ratio and by being dominated by various Arctostaphylos species as opposed to the interior dominant, Adenostoma fasciculatum. The maritime climate influence along the California central coast is associated with patterns of woody plant composition and β‐diversity among sites. Summer fog in coastal lowlands and higher winter precipitation in coastal uplands combine to lower late dry season water deficit in coastal chaparral and contribute to longer fire return intervals that are associated with obligate seeders and more local endemism. Soil nutrients are comparatively less important in explaining plant community composition, but heterogeneous azonal soils contribute to local endemism and promote isolated chaparral patches within the dominant forest vegetation along the coast.     

Vulnerability to xylem cavitation and the distribution of Sonoran Desert vegetation

We studied 15 riparian and upland Sonoran desert species to evaluate how the limitation of xylem pressure (Ψ(x)) by cavitation corresponded with plant distribution along a moisture gradient. Riparian species were obligate riparian trees (Fraxinus velutina, Populus fremontii, and Salix gooddingii), native shrubs (Baccharis spp.), and an exotic shrub (Tamarix ramosissima). Upland species were evergreen (Juniperus monosperma, Larrea tridentata), drought-deciduous (Ambrosia dumosa, Encelia farinosa, Fouquieria splendens, Cercidium microphyllum), and winter-deciduous (Acacia spp., Prosopis velutina) trees and shrubs. For each species, we measured the "vulnerability curve" of stem xylem, which shows the decrease in hydraulic conductance from cavitation as a function of Ψ(x) and the Ψ(crit) representing the pressure at complete loss of transport. We also measured minimum in situ Ψ(x)(Ψ(xmin)) during the summer drought. Species in desert upland sites were uniformly less vulnerable to cavitation and exhibited lower Ψ(xmin) than riparian species. Values of Ψ(crit) were correlated with minimum Ψ(x). Safety margins (Ψ(xmin)-Ψ(crit)) tended to increase with decreasing Ψ(xmin) and were small enough that the relatively vulnerable riparian species could not have conducted water at the Ψ(x) experienced in upland habitats (-4 to -10 MPa). Maintenance of positive safety margins in riparian and upland habitats was associated with minimal to no increase in stem cavitation during the summer drought. The absence of less vulnerable species from the riparian zone may have resulted in part from a weak but significant trade-off between decreasing vulnerability to cavitation and conducting efficiency. These data suggest that cavitation vulnerability limits plant distribution by defining maximum drought tolerance across habitats and influencing competitive ability of drought tolerant species in mesic habitats.     

Ecological and evolutionary differences between Mediterranean seeders and resprouters

Abstract. Differences in allocation patterns between seeders and resprouters in several Mediterranean plant communities (Australia, California and South Africa) have led to the prediction that seedlings of seeders grow faster than those of resprouters. In the Mediterranean Basin, it has also been hypothesized that regeneration strategy of plants after fire is associated with several other life history traits. This paper tests both hypotheses for the dominant plants in the Mediterranean Basin from literature data. Results show that seeders from the Mediterranean Basin grow significantly faster and allocate more biomass to leaf plus paracotyledons than resprouters. Seeders are mainly non‐sclerophyllous, anemochorous, dry‐fruited, small‐seeded species that evolved in the Quaternary (post‐Pliocene) and are associated with earlier successional stages. Resprouters are mainly sclero‐phyllous, vertebrate‐dispersed, fleshy‐fruited, large‐seeded species that evolved in the Tertiary (pre‐Pliocene) and are associated with late successional stages.     

Recovery patterns of three chaparral shrub species after wildfire

Summary In a mature, even aged stand of mixed chaparral, Rhus laurina (facultative resprouter) had consistently higher water potentials and deeper roots than Ceanothus spinosus (facultative resprouter) and Ceanothus megacarpus (obligate seeder). For two years following a wildfire, the same stand of chaparral had resprouts with higher survivorships, predawn water potentials, stomatal conductances, photosynthetic rates and shoot elongation rates than seedlings. Supplemental irrigation of seedlings during summer months removed differences between resprouts and seedlings suggesting that the cause of such differences was limited water availability to the shoot tissues of seedlings. After two years of postfire regrowth, mean seedling survivorship for the obligate seeder (C. megacarpus) was 42%, whereas seedling survivorship for facultative resprouters was only 18% (C. spinosus) and 0.01% (R. laurina). Our results are consistent with the hypothesis that lack of resprouting ability among obligate seeders is offset by an enhanced ability to establish seedlings after wildfire, allowing obligate seeders to maintain themselves in mixed populations through many fire cycles.     

SURVIVAL OF SEVERE DROUGHT BY A NON-SPROUTING CHAPARRAL SHRUB

A 2-yr drought (1975-1977) in much of California provided unusually severe water stress on many native plant species. In this paper we evaluate the effects of this drought on Arctostaphylos viscida Parry, a common non-sprouting chaparral shrub in the foothills of the southern Sierra Nevada. At the peak of the drought, water potentials as low as –74 bars were measured. While only three of 90 shrubs sampled were killed by the drought, all but one showed signs of drought-induced dieback. First and second year post-drought twig growth was significantly greater on shrubs with 90% or greater branch mortality than on those suffering lesser dieback. In all cases, new growth occurred only on surviving branches. Midday water stress measurements showed little difference for shrubs exhibiting high and low levels of dieback. These results suggest that drought survival in Arctostaphylos viscida may depend on a mechanism where certain branches are sacrificed with the surviving ones preferentially receiving available resources. The possibility that this may involve the dieback of below ground roots and/or lack of production of new roots to compensate for lack of root turnover is supported by water stress measurements of adjacent fire damaged shrubs. Implications of possible specialized drought survival mechanisms in non-sprouting shrubs are explored.     

Do xylem fibers affect vessel cavitation resistance?

Possible mechanical and hydraulic costs to increased cavitation resistance were examined among six co-occurring species of chaparral shrubs in southern California. We measured cavitation resistance (xylem pressure at 50% loss of hydraulic conductivity), seasonal low pressure potential (P(min)), xylem conductive efficiency (specific conductivity), mechanical strength of stems (modulus of elasticity and modulus of rupture), and xylem density. At the cellular level, we measured vessel and fiber wall thickness and lumen diameter, transverse fiber wall and total lumen area, and estimated vessel implosion resistance using (t/b)(h)(2), where t is the thickness of adjoining vessel walls and b is the vessel lumen diameter. Increased cavitation resistance was correlated with increased mechanical strength (r(2) = 0.74 and 0.76 for modulus of elasticity and modulus of rupture, respectively), xylem density (r(2) = 0.88), and P(min) (r(2) = 0.96). In contrast, cavitation resistance and P(min) were not correlated with decreased specific conductivity, suggesting no tradeoff between these traits. At the cellular level, increased cavitation resistance was correlated with increased (t/b)(h)(2) (r(2) = 0.95), increased transverse fiber wall area (r(2) = 0.89), and decreased fiber lumen area (r(2) = 0.76). To our knowledge, the correlation between cavitation resistance and fiber wall area has not been shown previously and suggests a mechanical role for fibers in cavitation resistance. Fiber efficacy in prevention of vessel implosion, defined as inward bending or collapse of vessels, is discussed.     

The impact of the invasive alien grass Cortaderia jubata (Lemoine) Stapf on an endangered mediterranean-type shrubland in California

Abstract. Since its introduction in the late 1800s, the perennial tussock grass Cortaderia jubata (Lemoine) Stapf has become an increasingly frequent member of coastal plant communities in California and Oregon. In this study, the community changes associated with C. jubata invasion into the mediterranean-type vegetation of Vandenberg Air Force Base, California were examined. Pristine plots of maritime chaparral were compared with spatially and topographically matched plots dominated by C. jubata . Aerial photographs indicated that the invaded plots had previously been shrubland. C. jubata invasion created a structurally less complex perennial grassland that was markedly depauperate in native shrub species. The absence of native shrubs depressed native richness in jubata grassland, but the greater richness of both native and alien herbaceous species made overall richness in jubata grassland indistinguishable from maritime chaparral. Vegetational differences were associated with differences in arthropod and small mammal populations between vegetation types. Arthropod abundance and order diversity were lower in plots dominated by C. jubata than in adjacent chaparral. Insect traps in C. jubata plots contained a significantly smaller proportion of Hymenoptera and Homoptera and a significantly greater proportion of Aràneae than traps in maritime chaparral. Rodent activity was significantly lower while rabbit activity was significantly greater in jubata grassland compared to maritime chaparral. This study indicates that the presence of C. jubata can dramatically alter the mediterranean-type landscape of central California. While it is likely that the initial establishment of C. jubata is associated with disturbances that are common in this ecosystem, the ability of C. jubata to expand from founding populations and to persist for long periods of time pose a serious threat to the native diversity of these unique systems.     

Heavy browsing affects the hydraulic capacity of Ceanothus rigidus (Rhamnaceae)

Defoliation by herbivores can reduce carbon assimilation, change plant water relations, and even shift the biotic structure of plant communities. In this study, we took advantage of a long-term deer exclosure experiment to examine the consequences of persistent deer herbivory on plant water relations and the xylem structure–function relationships in Ceanothus rigidus, a maritime chaparral shrub in coastal California. Browsed plants had thicker stems with many intertwined short distal twigs, and significantly higher sapwood-to-leaf area ratios than their non-browsed counterparts. Leaf area-specific hydraulic conductivity was similar in both browsed and non-browsed plants, but xylem area-specific conductivity was significantly lower in the browsed plants. Vessel diameters were equivalent in both plant groups, but the number of vessels on a transverse area basis was nearly 40 % lower in the browsed plants, accounting for their lower transport efficiency. Mid-day in situ water potentials and losses of hydraulic conductivity due to embolism were similar in both groups of plants but stomatal conductance was higher in the browsed shrubs in the early part of the growing season. We discuss our findings in the context of whole-plant ecophysiology, and explore the consequences of herbivory on hormonal signals, wood anatomy, and xylem function.     

WATER RELATIONS OF TWO CALIFORNIA CHAPARRAL SHRUBS

Water use patterns of two California chaparral shrub species, chamise (Adenostoma fasciculatum H. and A.) and Stanford manzanita (Arctostaphylos stanfordiana Parry), were compared during summer drought. Observations of diurnal and seasonal courses of shoot water potential, leaf conductance and transpiration revealed that chamise was more conservative in water use than manzanita. Evidence obtained cast doubt on a hypothesis previously proposed to explain an anomalous pattern of shoot water potential in chamise.     

Woody species tolerance to expansion of the perennial tussock grass Ampelodesmos mauritanica after fire

Abstract. In Mediterranean shrublands, post‐fire accumulation of above‐ground biomass of resprouters is faster than that of seeders. This suggests that resprouters may have a competitive advantage. To test this hypothesis, we used a removal experiment to study the effect of the presence of the dominant tussock‐grass Ampelodesmos mauritanica on the resprouting shrubs Erica multiflora and Globularia alypum and on the seeders Rosmarinus officinalis and Pinus halepensis three and four years after a wildfire. Water potential of target plants was also measured to see if Ampelodesmos removal increased water availability. Ampelodesmos marginally reduced growth of all target species but did not influence survival or water potential of any target species. Our results suggest that the effect of climatically influenced water stress was stronger than the effect of Ampelodesmos neighbours. Plant‐plant interactions in this Mediterranean community are weak after fire and the magnitude of the Ampelodesmos effect does not differ between seeders and resprouters.     

Root systems of chaparral shrubs

Summary Root systems of chaparral shrubs were excavated from a 70 m² plot of a mixed chaparral stand located on a north-facing slope in San Diego County (32°54 N; 900 m above sea level). The main shrub species present were Adenostoma fasciculatum, Arctostaphylos pungens, Ceanothus greggii, Erigonum fasciculatum, and Haplopappus pinifolius. Shrubs were wired into their positions, and the soil was washed out beneath them down to a depth of approximately 60 cm, where impenetrable granite impeded further washing and root growth was severely restricted. Spacing and interweaving of root systems were recorded by an in-scale drawing. The roots were harvested in accordance to their depths, separated into diameter size classes for each species, and their dry weights measured. Roots of shrubs were largely confined to the upper soil levels. The roots of Eriogonum fasciculatum were concentrated in the upper soil layer. Roots of Adenostoma fasciculatum tended to be more superficial than those from Ceanothus greggii. It is hypothesized that the shallow soil at the excavation site impeded a clear depth zonation of the different root systems. The average dry weight root:shoot ratio was 0.6, ranging for the individual shrubs from 0.8 to 0.4. The root area always exceeded the shoot area, with the corresponding ratios ranging from 6 for Arctostaphylos pungens to 40 for Haplopappus pinifolius. The fine root density of 64 g dry weight per m² under the canopy was significantly higher than in the unshaded area. However, the corresponding value of 45 g dry weight per m² for the open ground is still high enough to make the establishment of other shrubs difficult.     

Tissue water relations of three chaparral shrub species after wildfire

Summary We compared the tissue water relations among resprouts and seedlings of three chaparral species during the first summer drought after wildfire. Two of the species, Rhus laurina and Ceanothus spinosus recover after fire by a combination of resprouting and seedling establishment (facultative resprouters), whereas a third species, Ceanothus megacarpus recovers by seedling establishment alone (obligate seeder). Our objectives were to document any differences in tissue water characteristics that might arise between resprouts and seedlings and to test the hypothesis that seedlings of obligate seeders develop more drought tolerant characteristics of their tissues than seedlings of facultative resprouters. We found that resprouts had much higher predawn values of water potential, osmotic potential, and turgor potentials than seedlings. Predawn turgor potentials of resprouts were 1.5 MPa through July and August when turgor potentials for seedlings remained near 0 MPa. During summer months, midday water potentials were 2 to 3 MPa higher for resprouts than seedlings and midday conductances of resprouts were two to five fold greater than those of seedlings. Even though resprouts did not experience severe water stress like seedlings, their tissue water characteristics, as determined by pressure-volume curve analyses, were similar by the peak of the drought in August. Further-more, the tissue water characteristics of seedlings from the obligate seeder, C. megacarpus, were similar to those of facultative resprouters — R. laurina, and C. spinosus. We attribute the observed differences in plant water status between resprouts and seedlings to differences in rooting depths and access to soil moisture reserves during summer drought. We conclude that the higher growth rates, photosynthetic performance, and survivorship of postfire resprouts are primarily a result of higher water availability to resprouting tissues during summer months. It appears that the greater seedling survivorship during summer drought observed for the obligate seeder, C. megacarpus, is not associated with more favorable tissue water characteristics.     

Water stress tolerance of shrubs in Mediterranean-type climate regions: Convergence of fynbos and succulent karoo communities with California shrub communities

Mediterranean-type climate regions are highly biodiverse and predicted to be particularly sensitive to climate change. Shrubs of the mediterranean-type climate region of South Africa are highly threatened, and their response to water stress has been comparatively little studied. Resistance to water stress induced xylem cavitation (P(50)) and xylem specific hydraulic conductivity (K(s)) were measured in 15 shrub species from fynbos and succulent karoo communities of South Africa. Species displayed a fivefold variation in cavitation resistance (P(50) of -1.9 to -10.3 MPa) with succulent karoo species displaying greater interspecific variability in P(50) than fynbos species. Principal components analysis (including P(50), minimum seasonal water potential, K(s), and xylem density) showed the response to water stress in fynbos species to be similar to chaparral species from the mediterranean-type climate region of California. The data suggest convergence of community and species-specific water stress "strategies" between these mediterranean-type climate regions with respect to their xylem traits. On the basis of the current study and reported plant death and dieback in these regions, woody species within the fynbos may be more susceptible to climate warming and drying than those within the succulent karoo that appear to be utilizing more diverse xylem strategies in response to water stress.     

Cavitation resistance and seasonal hydraulics differ among three arid Californian plant communities

Vulnerability to water stress-induced cavitation was measured on 27 woody shrub species from three arid plant communities including chaparral, coastal sage and Mojave Desert scrub. Dry season native embolism and pre-dawn water potential, and both wet and dry season xylem specific hydraulic conductivity (Ks) were measured. Cavitation resistance, estimated as water potential at 50% loss in conductivity (Psi50), was measured on all species during the wet season and on a subset of species during the dry season. Cavitation resistance varied with sampling season, with 8 of 13 sampled species displaying significant seasonal shifts. Native embolism and water potential were useful in identification of species displaying seasonal shifts. The Ks was not different among sites or seasons. The Psi50 varied among species and communities. Within communities, interspecific variation may be partially explained by differences in rooting depth or leaf habit (evergreen, semi-deciduous, deciduous). Communities diverged in their Psi50 with chaparral species displaying the greatest cavitation resistance regardless of sampling season. The greater cavitation resistance of chaparral species is surprising, considering the greater aridity of the Mojave Desert site. Adaptation to arid environments is due to many plant traits, and aridity does not necessarily lead to convergence in cavitation resistance.     

Tissue water relations of four co-occurring chaparral shrubs

Summary Chaparral shrubs of California have a suite of morphological and physiological adaptations to withstand the prolonged summer droughts of a mediterranean climate. Not all species of chaparral have the same rooting depth and there is some evidence that those with shallow roots have tissue that is most tolerant to water stress. We tested this notion by comparing the tissue water relations of four co-occurring chaparral shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a pressure-volume technique and a dew-point hygrometer to metsure seasonal changes in osmotic potential when plant tissue was fully hydrated and osmotic potential at predawn, midday, and the turgor loss point. We also calculated seasonal changes in the minimum daily turgor potential, saturated weight/dry weight ratio of leaf tissue, and the bulk modulus of elasticity. We had information on the seasonal water use patterns and apparent rooting depths of these same four shrubs from a previous study (Davis and Mooney 1986). All evidence indicated that Rhamnus had shallow roots and Quercus deep roots. Our results indicated that the tissue water relations of our four co-occurring chaparral shrubs were not alike. Even though Rhamnus had shallow roots, it had the least xerophytic tissue. Seasonal osmotic potential and saturated weight/dry weight ratios were relatively high and bulk modulus of elasticity and minimum daily turgor potentials were low. Furthermore, even though Quercus had deep roots and experienced no seasonal water stress at our study site, its tissue water relations indicated relatively high tolerance to water stress. We conclude that seasonal drought tolerance of stem and leaf tissue of co-occurring chaparral shrubs does not necessarily correspond to rooting depth, to soil moisture resources available to the shrub, or to the degree of seasonal water stress experienced by the shrub.     

Comparative community physiology: nonconvergence in water relations among three semi-arid shrub communities

Plant adaptations to the environment are limited, and therefore plants in similar environments may display similar functional and physiological traits, a pattern termed functional convergence. Evidence was examined for functional convergence among 28 evergreen woody shrubs from three plant communities of the semi-arid winter rainfall region of southern California. Both leaf and water relations traits were examined, including seasonal stomatal conductance (gs), specific leaf area (SLA), leaf specific conductivity (Kl), seasonal water potential (Psi w), stem cavitation resistance (Psi 50), and xylem density. Species display community-specific suites of xylem and leaf traits consistent with different patterns of water use among communities, with coastal sage scrub species utilizing shallow pulses of water, Mojave Desert scrub species relying on deeper water reserves, and chaparral species utilizing both shallow and deep moisture reserves. Communities displayed similar degrees of water stress, with a community-level minimum Psi w (Psi wmin) of c. -4.6 Mpa, similar to other arid communities. Pooled across sites, there was a strong correlation between Psi wmin and xylem density, suggesting that these traits are broadly related and predictive of one another. This comparative community physiology approach may be useful in testing hypotheses of functional convergence across structurally similar semi-arid communities.     

GROWTH FORMS IN ARCTOSTAPHYLOS GLAUCA

Arctostaphylos glauca has evolved two growth forms in the San Bernardino Mountains of Southern California. A chaparral ecotype of upright shrubs with relatively long leaves occurs in chaparral on the south and west sides of the mountain range. A desert woodland ecotype of low, compact, spreading shrubs with shorter leaves occurs in pinyon woodland on the north and northeast sides of the same range. The chaparral ecotype flowers early and fruits heavily. Upon periodic destruction by fire its populations are replaced by numerous seedlings. The woodland ecotype flowers late and fruits sparingly. Its lower seed reproductive potential is further indicated by the lower total number of floral units, by a lower proportion of fruit set, and by an apparent truncation of the flowering season before all the flower buds open. An alternative strategy, involving long life, compact growth form, and the exploitation of new soil resources by layering and vegetative reproduction in the absence of regular fire recycling has evidently adapted an old chaparral plant to recent desert woodland conditions.     

Comparative water relations of adjacent california shrub and grassland communities

Summary Much of the coastal mountains and foothills of central and southern California are covered by a mosaic of grassland, coastal sage scrub, and evergreen sclerophyllous shrubs (chaparral). In many cases, the borders between adjacent plant communities are stable. The cause of this stability is unknown. The purpose of our study was to examine the water use patterns of representative grasses, herbs, and shrubs across a grassland/chaparrel ecotone and determine the extent to which patterns of water use contribute to ecotone stability. In addition, we examined the effects of seed dispersal and animal herbivory. We found during spring months, when water was not limited, grassland species had a much higher leaf conductance to water vapor diffusion than chaparral plants. As the summer drought progressed, grassland species depleted available soil moisture first, bare zone plants second, and chaparral third, with one chaparral species (Quercus durata) showing no evidence of water stress. Soil moisture depletion patterns with depth and time corresponded to plant water status and root depth. Rabbit herbivory was highest in the chaparral and bare zone as indicated by high densities of rabbit pellets. Dispersal of grassland seeds into the chaparral and bare zone was low. Our results support the hypothesis that grassland species deplete soil moisture in the upper soil horizon early in the drought, preventing the establishment of chaparral seedlings or bare zone herbs. Also, grassland plants are prevented from invading the chaparral because of low seed dispersability and high animal herbivory in these regions.     

Legume seeders of the Cape Floristic Region inhabit more fertile soils than congeneric resprouters—sometimes

In fire-prone ecosystems, plants for the most part persist via either soil-stored seed banks (seeders) or below-ground storage structures (resprouters). Given their greater allocation of resources above ground to growth and reproduction, seeders are likely to have a higher nutrient requirement than resprouters. This may result in discernable differences in habitat nutrition and leaf morphology. These differences are probably accentuated in Cape legumes given their poor adaptation to low-P soils. It was hypothesized that legume seeders occupy habitats with greater fertility and possess larger, less sclerophyllous leaves than resprouters. Site nutrition and leaf morphologies were compared between seeders and resprouters in the genera Otholobium and Psoralea. There were no differences in leaf morphology between seeders and resprouters. Seeders sites had a higher total [N], exchangeable [Ca] and [Mg], and CEC, but lower [Fe] than resprouters. Only within Otholobium, did seeder sites have a higher Bray II [P]. This genus-specific variation in available P is probably a consequence of greater variation in soil type and precipitation between seeders and resprouters. Conversely, niche construction may contribute to the differences in soil fertility between seeders and resprouters in Psoralea. Thus, our data showed a general tendency for seeders to inhabit more fertile sites than resprouters. Caution is required, however, in generalizing these results, as our data indicate a difference in factors affecting soil nutrient availability between legume genera. Changes in soil fertility post-fire may limit legume persistence beyond the early stages of succession.