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 California's 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.     

Plant community response following the removal of the invasive Lupinus arboreus in a coastal dune system

The removal of invasive species is common in restoration projects, yet the long‐term effects of pest management programs are seldom assessed. We present results of a long‐term program to remove the invasive species Lupinus arboreus (lupin) from sand dunes in New Zealand. We evaluate the response of plant communities to lupin removal, by comparing total plant cover, the cover of non‐native and native plant species, and species richness between sand dune sites where lupin removal has occurred, not occurred, and where lupin has never been present. Neither lupin presence nor removal had a significant impact on the foredune environment. Following removal, total and other non‐native plant cover remained higher, and the cover of several native sand dune species remained lower compared with uninvaded sites in the deflation and backdune environments. These changes can be attributed to persistent effects associated with the invasion of lupin, but have also developed in response to lupin removal. The results of this study have implications for restoration projects in sand dunes. Pest management alone is unlikely to be sufficient to restore plant communities. Given the difficulties in restoring plant communities once an invasive species has established, managers should prioritize actions to prevent the spread of invasive species into uninvaded areas of sand dunes. Finally, the response to lupin invasion and removal differed between dune habitats. This highlights the importance of tailoring a pest management program to restoration goals by, for example, prioritizing areas in which the impacts of the invading species are greatest.     

Promotion of mid-successional seedling recruitment and establishment by Juniperus virginiana in a coastal environment

Positive interactions between Juniperus virginianaand woody seedlings may influence trends in primary succession along theAtlantic Coast of eastern North America. Woody species richness was greaterbeneath isolated J. virginiana trees than in more exposed,grass-covered dune sites on a Virginia, USA barrier island. Fleshy fruitedseedswere more abundant in the seed bank beneath J. virginianathan in adjacent exposed sites, suggesting that J.virginiana may be utilized by passerine birds. Photosyntheticallyactive radiation was reduced below J. virginiana, and soiltemperature fluctuations were moderated during the growing season. In addition,moisture content, organic matter, and chlorides were higher for soils underJ. virginiana than in exposed sites. For plantedPrunus serotina and Sassafras albidumseedlings, mortality was lower beneath J. virginiana ascompared with the exposed treatment; however, no S.albidumseedlings survived beyond mid-August. The effect of J.virginiana on the recruitment and distribution of mid-successionalwoody seedlings in coastal environments may be passive, through the non-randomdistribution of fleshy seeds by perching birds, or active, through increasedseedling survival due to J. virginiana initiatedalterations in microclimate and edaphic factors.     

The benthic macroinvertebrate community in a coastal sand dune lake relative to habitat and changing lake levels

The benthic macroinvertebrate community (BMI) in a freshwater coastal dune lake without a surface outlet was investigated in May and October, 1986. Fifty-three invertebrate taxa were identified from Carter Lake, including three euryhaline crustacean species (Corophium spinicorne, Gnorimosphaeroma oregonensis lutea, and Acanthomysis awatchensis). Corophium spinicorne dominated the BMI communities of the littoral zones and sphaeriid clams dominated the deepwater community.The lake level dropped about 2.5 m between April and October. Based upon this decline, the lake bottom was divided into four major habitats: a sandy temporarily submerged littoral zone (A); a sandy submerged littoral zone (B); and mid-depth zone of mixed mud and sand and the macrophyte, Nitella (C); and a deep zone (D) with soft mud. The average density of BMI was highest in the littoral zones (A and B) in May and in zone B in October (zone A was dry). The lowest density occurred in zone D. In May, BMI biomass was highest in the littoral zones, but the biomass was highest in the mid-depth zone in October. The mid-depth zone in October. The mid-depth zone had the most diverse community.The two most abundant species in the temporarily submerged area, Corophium spinicorne and Juga plicifera, were found in greater numbers deeper in the lake after the water level dropped, suggesting migration by these species in response to changing water levels.     

Influence of woody invader control methods and seed availability on native and invasive species establishment in a Hawaiian forest

When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai’i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but short-lived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.     

Sandblasting as a possible factor controlling the distribution of plants on a coastal dune system

Intensity of the abrasive effect of wind-borne sand – sandblasting – in addition to other environmental factors was measured at two vegetation zones on a sandy beach and one site at an inland area. One zone on the beach included foredunes sparsely vegetated by dune species such as Carex kobomugi and Calystegia soldanella. The other zone which was located ∼50 m inland from the first zone was flat grassland dominated by inland species such as Miscanthus sinensis and Imperata cylindrica var. Koenigii. The inland site consisted of short grassland located 3 km inland from the beach. Intensity of sandblasting was estimated by the whiteness of a transparent plastic sheet exposed to the air for 2 weeks. This sheet turned whitely opaque when it was abraded by wind-borne sand. The other environmental factors measured at the beach were intensity of salt spray, soil water content, soil salinity, and sand accumulation, while intensity of salt spray was the only additional factor measured at the inland site. Intensity of sandblasting was considerably higher at the foredune zone, while that at the grassland zone was as low as that at the inland site. Considerable salt spray was detected at the foredune and grassland zones. Differences in other environmental factors were small between the two zones on the beach. In order to compare the difference in tolerance to sandblasting, a jet of sand was applied to one ordinary species, C. kobomugi, from the foredune and two species, M. sinensis and I. cylindrica, from the grassland zone. The difference in tolerance was determined by the decrease in the area of green leaf after applying sandblasting with commercial sandblaster and/or spraying with sea water. M. sinensis and I. cylindrica lost much of the leaf area after sandblasting and salt spraying, while C. kobomugi lost little. These results indicated that one of the characteristic environmental factors of a foredune is the high intensity of sandblasting accompanied by salt spray, and that species found in the foredune are more tolerant to sandblasting than species distributing in more inland areas.     

Carbon addition interacts with water availability to reduce invasive forb establishment in a semi-arid grassland

Increases in nitrogen (N) availability can favor fast-growing invasive species over slow-growing native species. One way to reduce N availability is to add labile carbon (C) to the soil, which can lead to microbial immobilization of plant available N. This method has been used, with widely varying degrees of success, to both study and control plant invasions. One reason that C addition might not work as expected is that N is not always the limiting resource for plant growth. For example, if plant growth is limited by water, changes in N availability might have little effect on invasion. Here I ask whether effects of C addition on N availability, resident plant biomass, and invasion depend on water availability in semi-arid mixedgrass prairie. Six invasive species were seeded into plots treated with a factorial combination of water (ambient or added) and N (+C, control or +N). Carbon addition reduced capture of mineral N by resin probes (by an average of 73%), and reduced biomass of resident species (from 336 g m⁻² to 203 g m⁻²), both with and without added water. In contrast, because there was little invasion in ambient-water plots, C addition reduced invasion only in added-water plots. Given added water, C addition reduced biomass of Centaurea diffusa by 95%, and prevented invasion by Gypsophila paniculata and Linaria dalmatica. Mechanisms by which C addition reduced invasion varied by species, with added C reducing the growth of individual C. diffusa plants, but reducing numbers of G. paniculata and L. dalmatica individuals.     

Lizard herbivory in a sand dune environment: the diet of Angolosaurus skoogi

Summary Early observations of the feeding habits of Angolosaurus skoogi (Andersson), a 50–120 g lizard inhabiting the almost vegetationless dunes of the northern Namib Desert, indicated that this species was herbivorous. Less than two percent of living lizard species are predominantly herbivorous and these species tend to have a high body mass (>100 g). We therefore analysed fecal pellets of A. skoogi to obtain dietary details of this relatively rare lizard. Eighteen food taxa were identified: 81 percent by mass was composed of vegetable matter, of which 56 percent was an endemic, perennial cucurbit Acanthosicyos horrida. Both spatial and temporal variation in dietary composition were apparent. Ready access to water in the form of A. horrida is postulated to have allowed Angolosaurus to occupy this extreme habitat and to forage on components of the windblown detritus, a food source that would probably not be usable in other circumstances.     

The effect of stolon fragmentation on the colonization of clonal invasive Carpobrotus edulis in a coastal dune system: a field test

Disturbances usually initiate processes of fragmentation in clonal plants, with the consequent division into portions of different size. The ability of these portions to survive and regrow after fragmentation plays an important role in the maintenance of populations and the colonization of new environments. In this field experiment we aim to determine the importance of stolons as reserve organs in the colonization of a coastal sand dune by a clonal invader. We simulated an event of fragmentation of clones of an aggressive invader into portions with short and long stolon sizes. Our results showed a reduction of biomass allocation to roots in the long stolon treatment that was balanced by an increase in the above‐ground growth; consequently, the area colonized by the invader was greater. We report evidence that stolons can contribute to buffering stressful conditions and allow expansion of the invader into a natural coastal sand dune.     

Long term dynamics of Rosa multiflora in a successional system

Long term studies of invasion dynamics are critical in developing a more complete understanding of the factors that influence species spread. To address this issue, the dynamics of the non-native invasive plant, Rosa multiflora, were examined using a 40-year record of successional change. The roles of biotic and abiotic factors in regulating R. multiflora invasion were also assessed. The invasion showed an initial 9-year time lag, followed by a 20-year period of population expansion and an ultimate decline as succession progressed. During all phases of R. multiflora’s invasion, there was continuous turnover within plots. Rainfall during the previous season was found to increase R. multiflora colonization during population expansion while tree species inhibited the invader’s growth. During expansion and decline of R. multiflora, common associated species were often positively or negatively correlated with changes in R. multiflora cover. Though early population dynamics were regulated by propagule pressure, the major influence on R. multiflora late in succession was canopy closure. Although the invasion of this species was largely self-limiting in this system, the species is likely to persist within late successional systems and may require management intervention.     

Germination responses to temperature responsible for the seedling emergence seasonality ofPrimula sieboldii E. Morren in its natural habitat

Temperature requirements for the breaking of seed dormancy and germination inPrimula sieboldii E. Morren and the annual surface-soil temperature regime in one of its natural habitats were investigated in order to clarify the germination responses determining the seedling emergence seasonality of the species. In a grassland nature reserve in an abandoned flood plain of the Arakawa River, natural seedling emergence of the species was shown to be restricted to mid- to late-spring before the closure of seasonal vegetational gaps, when the daily mean soil surface temperature reached about 15°C, accompanied by large daily fluctuations of about 10°C. Mature seeds collected in late June were never able to germinate at any constant temperature in the range of 8–40°C unless they had been previously subjected to moist-chilling treatment. The proportion of seeds which were released from dormancy increased with increasing duration of the moist-chilling treatment at 2°C, 70–85% of seeds becoming germinable at 16–28°C after 12 weeks of pretreatment at 2°C. The thermal time required for the germination of the thus-pretreated seed population was 905–1690 Kh with a base temperature of around 5°C. Fluctuating temperatures between 24°C and 16 or 12°C had a remarkable dormancy-breaking effect, inducing considerably quick germination in most of the seeds previously subjected to 2°C moist-chilling for 8 weeks.     

胜红蓟种子萌发/出苗对环境因子的响应

胜红蓟为我国南部低海拔区域农田和果园恶性杂草。为揭示胜红蓟环境适应规律、明确其危害性,评价和分析了胜红蓟种子萌发对光照、温度、pH、盐度、渗透势等环境因子的响应特征。结果表明:胜红蓟种子在10—30℃内均能萌发,最适萌发温度为20℃,交替温度处理可提高种子的萌发率。胜红蓟种子萌发对光照敏感,无光照处理种子不萌发。土壤相对湿度在50%—100%范围内,种子出苗率均高于55%,最适相对湿度为70%。胜红蓟种子仅能在土壤表面萌发,1 cm的播种深度完全抑制种子萌发。pH值在5—10之间,种子萌发率高于85%,最适pH值为7。NaCl浓度在0—80 mmol/L以内种子萌发率超过88%,浓度为160 mmol/L时萌发率接近30%。渗透势在-0.40—0 MPa以内种子萌发率高于70%,渗透势为-0.6 MPa时种子不能萌发。由此可见,胜红蓟种子萌发具有广阔的环境适生范围,对农作物具有较强的早期危害性。     

Sowing and transplantation of Adonis ramosa (Ranunculaceae) for the establishment of a new population in a seminatural broad-leaved deciduous forest floor

Adonis ramosa (Ranunculaceae) is a plant whose risk of extinction is increasing in Japan. In this study, we show that there is a significant possibility of establishing an A. ramosa population by sowing seeds or transplanting plants in a seminatural broad-leaved deciduous forest floor. Immediately after seed collection in early June, seeds were sown either in pots that were watered periodically or in a deciduous forest floor where the soil surface was prepared. In the following spring, 30–40% of the seeds germinated. Of the one-year-old plants buried at a depth of 2 cm and two-year-old plants buried at a depth of 2 or 8 cm in the fall, 96% sprouted leaves in the following spring. However, 79% of the one-year-old plants buried at a depth of 8 cm sprouted leaves. In spring, the number of leaves, the widths of individual plants, and the heights of the two-year-old plants were greater than those of the one-year-old plants. The dry weights of one- and two-year-old plants in fall after one spring growing season increased by 1.6–2.3 times and by 2.3–5.1 times, respectively, compared to the dry weight at the time of transplanting. Also, the dry weights of the plants transplanted at a depth of 8 cm were higher than those of plants transplanted at a depth of 2 cm for both one- and two-year-old plants. The results suggest that the growth conditions at a depth of 8 cm are more favorable than those at a depth of 2 cm.     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

Seed viability and seed dormancy of non-native Phragmites australis in suburbanized and forested watersheds of the Chesapeake Bay, USA

The non-native, invasive haplotype of Phragmites australis is rapidly invading tidal and non-tidal wetlands across North America. Phragmites has the potential to spread by seeds and rhizomes. Seed viability and dormancy differences were quantified among 18 patches of non-native Phragmites in subestuarine wetlands in developed (i.e., suburbanized) vs. forested watersheds of the Chesapeake Bay. We used tetrazolium and germination assays to assess seed viability and compared germination percentages and rate of germination among fresh seeds, cold–moist treated seeds, and warm–dry treated seeds to evaluate seed dormancy. Seed viability was <1% in most patches but a few patches produced abundant viable seeds (5–21%). Seed viability, however, did not differ significantly between wetlands in forested vs. developed watersheds. Contrary to studies of Phragmites seed dormancy in European populations, some Phragmites seeds were dormant at maturity; cold–moist treated seeds germinated faster and to higher percentages than fresh seeds or warm–dry treated seeds.     

Comparative population ecology of dune slack species: the relation between population stability and germination behaviour in brackish environments

Populations of short-lived pioneer species from brackish primary dune slacks and beach plains usually have to cope with unpredictably fluctuating abiotic conditions. Life history parameters play a major role in adaptation to temporal environmental heterogeneity. A population's susceptibility to fluctuating salinity levels may be largely determined by seed dormancy characteristics (e.g. seed variability, degree of enforced dormancy imposed by salinity) and juvenile salt resistance rather than by adult salt resistance.Nomenclature follows Hegi (1966).     

A comparison of sites suitable for the seedling establishment of two co-occurring species, Swintonia glauca and Stemonurus scorpioides, in a tropical peat swamp forest

Spatial and temporal ground-surface dynamics are major factors that affect regeneration and species coexistence in tropical peat swamp forests. We studied the seedling survivorship and morphological features of two tree species that play important roles in maintaining the ground-surface dynamics of a peat swamp forest in Sumatra. Large Swintonia glauca trees form mounds, whereas large Stemonurus scorpioides trees occupy non-mounds. We monitored the demography of naturally dispersed Swintonia and Stemonurus seedlings that germinated in 2000. Survivorship of Swintonia seedlings was high under conditions of late germination, high-light environment, and elevated ground surface, and was negatively affected by distance to the nearest conspecific adult. Survivorship of Stemonurus was high under conditions of early germination and high conspecific seedling density, and was also negatively affected by distance to the nearest conspecific adult. The allometric features of Stemonurus seedlings indicated characteristics of stress tolerance, that is, low growth rate and thick, porous roots. Stemonurus, which has large wingless seeds, regenerated in non-mounds around the parental trees, while winged Swintonia seeds dispersed farther from the parent and established in patchily distributed gaps and mounds. Thus, Swintonia seedlings can survive on non-mound sites within gaps and possibly create mounds, while Stemonurus seedlings tend to maintain non-mounds around the parental trees.     

Establishment and spread of founding populations of an invasive thistle: the role of competition and seed limitation

Successful plant invasions require both the founding and local spread of new populations. High plant densities occur only when founding plants are able to disperse their seeds well locally to quickly colonize and fill the new patch. We test this ability in a 7-year field experiment with Carduus acanthoides, an invasive weed in several North American ecosystems. Founder plants were planted in the center of 64 m² plots and we monitored the recruitment, distribution pattern, mortality, and seed production of the seedlings that originated from these founding plants. Competing vegetation was clipped not at all, once, or twice each year to evaluate the importance of interspecific competition. More seedlings recruited in the intermediate once-clipped plots, and these seedlings also survived better. The control plots had fewer microsites for seedling recruitment; clipping a second time in September stimulated grasses to fill up the gaps. The number of C. acanthoides recruits and their median distances from the founder plants were also explained by the initial seed production of the founding plants. Overall, the experiment shows that the success of founder plants can fluctuate strongly, as 55% of the plots were empty by the sixth year. Our study suggests that the local invasion speed following initial establishment depends strongly on both the propagule pressure and availability of suitable microsites for seedling recruitment and growth.     

Regional and local variation in seedling emergence,mortality and recruitment of a perennial herb in Mediterranean mountain habitats

Spatial patterns of seedling regeneration of woody Mediterranean plants are widely documented, with seedlings and juveniles growing almost exclusively under shrub and tree canopy. Neighbourhood habitat amelioration and consequent facilitative effects on recruitment have been extensively suggested as the major determinant of such spatial pattern for Mediterranean vegetation. Much less is known on recruitment patterns of perennial herbs. As herbs differ from woody plants in many relevant ecological aspects and in their life cycles, we would expect particularities in their recruitment dynamics. We analysed the spatial (regional, local and fine-scale) variation in environment (herb cover, litter depth, air temperature and irradiance) and its relationship with seedling emergence, mortality, and recruitment of the perennial herb Helleborus foetidus (Ranunculaceae) during 2 years. We conducted the study in three distant localities in south-eastern Iberian Peninsula (defining the regional scale). Local scale was defined in terms of microhabitat type based on three categories of vegetation cover (under evergreen shrubs or trees, under deciduous spiny-shrubs, and in open interspaces). Finally, fine-scale variation was defined in terms of 1 m² sampling plots. Our results showed regional, local and fine-scale variations in recruitment, with the major source of variation changing across scales. Seed input was important in determining differences in recruitment both at the largest (regional) and the finest scale. Environment had minor importance in shaping differences in recruitment at these two scales. At the local scale, variation between microhabitats was mainly related to differences in seedling survival through facilitative effects (alleviation of water and irradiance stresses) of shrub and tree cover, although such facilitative effect was not consistent in all localities or years. Our study thus points to a similarity in the spatial patterns of recruitment (and in the environmental factors shaping such patterns) between Mediterranean woody plants and perennial herbs, but much more effort is needed to assess the generality of such similarity.     

Influence of litter removal and mineral soil disturbance on the spread of an invasive grass in a Central Hardwood forest

Soil and litter disturbances within Central Hardwood forests may facilitate exotic plant species invasion of interior forest areas. Microstegium vimineum is an annual exotic grass that has become common throughout the Southeastern United States. Three replicates of three different mineral soil and litter disturbance treatments, plus a control with no disturbance, were established on the leading edge of M. vimineum patches prior to seed fall. All patches were located in areas with similar forest canopy structure and slope in three Central Hardwood forest stands prior to seed fall. At the beginning of the following growing season, each individual M. vimineum seedling was mapped within the treatment plots. The mean number of M. vimineum individuals that established within each treatment did not differ significantly from the control. The distance at which 90% of the individuals had spread, and the overall mean distance spread were significantly farther for the litter removal treatment than the control. The farthest individual seedling from the boundary of existing patches in both the litter removal and the mineral soil disturbance and litter removal treatments were significantly farther than the control. The individuals that spread the farthest are of most concern due to the large number of viable seed that a single M. vimineum plant can produce. These results suggest that disturbance of the litter layer may increase the spread rate of M. vimineum and facilitate its invasion of new habitats, and that leaving litter layers intact may slow the spread of M. vimineum.