Mark I. Borchert

Early postfire seed dispersal, seedling establishment and seedling mortality of Pinus coulteri (D. Don) in central coastal California, USA

Seedling recruitment in many highly serotinous populations of Pinus coulteri on Californias central coast depends almost entirely on periodic, stand-replacing fire. Compared to serotinous pines of the Mediterranean Basin, little detailed information is available on the postfire demography of California closed-cone pines, including P. coulteri. In September 1996 a wildfire burned the 760-ha American Canyon Research Natural Area (RNA). Using aerial photography, we mapped burn severity of P. coulteri-chaparral woodlands and forests within the RNA. From May to September of 1997, we also quantified seedling establishment and mortality in relation to biophysical site characteristics including fire severity. Seventy-six percent of P. coulteri forests and woodlands experienced high-severity burns, 9% moderate-severity burns, and 15% low-severity or unburned. Of the 53 plots used for seedling counts, 70% were high-severity, 26% moderate-severity, and 4% low-severity. Seedling densities 13 months postfire were low (0.21 m−2), but seedling mortality also was low (8.4%). Aerial seed bank size increased from north-facing to south-facing slopes and from high-severity to low-severity burns. Seedling recruitment was unrelated to burn severity and increased with the size of the canopy seed bank (cone density). Many seedlings established from rodent seed caches; 23% of the seedlings established in clumps from seeds cached by Dipodomys agilis, Chaetodipus californicus and Peromyscus maniculatus. Pinus coulteri seeds have low potential for dispersal by wind, but secondary dispersal by rodents moves seeds away from source trees and into neighboring chaparral. We discuss the potential importance of rodent seed caching to postfire demography of California and Mediterranean serotinous pines.

A comparative analysis of seed and cone characteristics and seed-dispersal strategies of three pines in the subsection Sabinianae

The seed-dispersal systems of Coulter pine (Pinus coulteri), gray pine (P. sabiniana), and Torrey pine (P. torreyana), all of the subsection Sabinianae, are not well understood. These pines occur in arid and semi-arid foothills and mountains of California that are subjected to frequent fires. Cone and seed traits of these three California pines are compared to those of four species of pines (sugar pine, P. lambertiana; Jeffrey pine, P. jeffreyi; ponderosa pine, P. ponderosa; and lodgepole pine, P. contorta) that occur in more mesic environments in the nearby Sierra Nevada mountains. The cones of the Sabinianae pines are large with thick, dense scales, and the scales of gray and Coulter pines are armed with sharp, recurved spines. The seeds of all three species are large, and those of gray and Torrey pines are nearly wingless. In contrast, the Sierra Nevada pines have small to medium-sized seeds with large wings that are initially dispersed by the wind. Heavy wing loading of the Sabinianae pine seeds causes them to fall rapidly, and they are not dispersed far by wind. However, animals remove the fallen seeds rapidly, and rodents and jays scatter hoarded many seeds in the soil. This caching activity results in seedling establishment. The unusual morphology of the cones and seeds of the Sabinianae pines is interpreted as a combination of traits that attract animal dispersers, thwart the foraging activities of seed predators, and promote the survival of seeds in an environment subject to frequent fires.

Reproduction and growth of the chaparral geophyte, Zigadenus fremontii (Liliaceae), in relation to fire

Zigadenusfremontii is often a striking component of the flora following firein the chaparral. Like other geophytes, it produces large numbers of flowers inthe first spring after a burn. Although these plants are most conspicuous intheearly postfire environment, the question that remains is, how do they persistinthe interval between fires? To address this we investigated differences in thegrowth and reproduction of Z. fremontii in burned andunburned chaparral. We monitored marked individuals for nine years at threesites: two that were burned in 1990 and one in the same area that was inunburned mature chaparral. We measured leaf area, and production of flowers andfruits. We also conducted seed experiments in the field to determine the ratesand timing of germination. We found that reproduction occurs only in theimmediate postfireperiod: flowering and production of fruits and seeds in the first yearfollowingfire, and seedling establishment by year 3. There was a cost of reproduction;plants that flowered (in the burn area) had negative growth rates the followingyear. In contrast, plants in unburned chaparral, which did not flower, hadpositive growth rates over the same period. Moreover, plants that produced themost flowers had the lowest growth rates. In the unburned chaparral site,plantswere not dormant as predicted from previous literature; instead they producedleaves nearly every year. In most years the average leaf area per plant wasgreater than that in the burned sites. Our results indicate that postfirereproduction depends on growth and carbohydrate storage in the inter-fireperiod. We also suggest that this species is relatively long-lived for aherbaceous perennial.

Patterns of post-fire flowering and fruiting in Chlorogalum pomeridianum var. pomeridianum (DC.) Kunth in southern California chaparral

For many geophytes living in Mediterranean ecosystems, the passage of fire can produce bursts of flowering, seed production, vegetative growth, and seedling recruitment. In the present study, we investigated patterns of flowering and fruit production of the chaparral geophyte Chlorogalum pomeridianum (common soap plant) at two sites: one burned in a prescribed fire and one in nearby unburned chaparral. Both sites burned in a wildfire 2 years later, and we continued monitoring marked plants for an additional 6 years, enabling us to observe the effects of recent reburning on reproduction and growth. We found that flowering was stimulated by fire but was not strictly fire-dependent. There was a positive relationship between bulb size and leaf area, as well as between these two characteristics and flower and fruit production. Flower stalk initiation occurred when plants reached a minimum leaf area of ~1000 cm2, indicating that a minimum bulb size must be reached before reproductive effort is initiated. Direct herbivory of flowering stalks reduced fruiting and leaf herbivory indirectly prevented the initiation of flowering stalks. In the first several years after fire, flower and fruit production could be explained by resource matching but in subsequent years, resource matching was replaced by resource switching.

Desiccation sensitivity and heat tolerance of Prunus ilicifolia seeds dispersed by American black bears ( Ursus americanus )

ABSTRACT. Carnivore consumption of fruit is a principle means by which many fleshy-fruited plant species achieve long-distance seed dispersal. We examined carnivore dispersal of hollyleaf cherry (Prunus ilicifolia) seeds, specifically assessing the survival, desiccation sensitivity, and germination of seeds found in bear scats. Studies were conducted both in the laboratory and in 2 burn areas in Los Padres National Forest, California. Bear scats containing P. ilicifolia seeds were collected in burned and unburned chaparral. We counted seeds in each scat and noted whether endocarps had tooth punctures or rattled audibly when shaken. For comparative germination trials, we also collected fruits and seeds directly from mature shrubs. In the laboratory, following a cold-moist stratification period, seeds were assessed for germinability. In the field, we compared desiccation rates and germinability of seeds from bear scats and freshly collected seeds. We compared rates of moisture loss and germination for seeds subjected to several different conditions, including 25 °C (room temperature), 30 °C and 65 °C (in the lab), and placement on exposed soil in a burn area (in the field), where midday temperatures were approximately 45 °C but likely much higher by late afternoon. Prunus ilicifolia seeds collected from bear scats were largely undamaged; the vast majority of these seeds germinated. In some cases, germination rate was higher for seeds from scats than for seeds from intact fruits. Several results indicate that desiccation is an important cause of reduced germinability. First, seeds that rattled audibly germinated poorly; and the louder the rattle, the lower the germination percentage. Second, seeds (both fresh and those from bear scats) placed in the field under protective screens had greatly reduced levels of germination (an 84% decline) after only 7 days. Third, seeds dried in the lab, even at relatively moderate temperatures, showed a decline in germination with seed moisture loss. The addition of high temperatures accelerated this decline in germination. We discuss the relevance of heat and desiccation sensitivity of seeds dispersed by bears to successful seed germination in burned and late-seral mesic and xeric chaparral.