V. Thomas Parker

Ecology of Soil Seed Banks

Examines factors influencing seed-bank dynamics and the variety of patterns found among different species. Topics include: the relationship of seed banks to vegetation dynamics; processes that influence inputs and losses from seed banks; and the role and importance of seed banks in vegetable types.

The New Paradigm in Ecology: Implications for Conservation Biology Above the Species Level

Conservation biology has traditionally focused on the fine scale and the species level of biological organization (Soule and Wilcox 1980), and biotic conservation is only one of the various goals that has directed the preservation and management of natural areas and resources. Resource management goals have largely been utilitarian or commercial, such as the maintenance of large areas for watersheds; preservation of marshes, bogs, and seasonal wetlands for flood water storage, nursery areas, and flow continuity in river systems, and the sustained yield of forests, fisheries, and wildlife (Temple et al. 1988; Aplet, Laven, and Fiedler in press). There have, however, been both practically and theoretically motivated calls for widening the focus to include scales beyond that of the individual site and levels of organization above that of the species (Soule 1989; Western 1989) to approach new goals. This essay will explore the relationship of this new frontier for conservation biology to advances in ecology. Much of the new ecological focus of conservation biology is driven by the shift in the overarching paradigm of ecology itself. We begin by defining the concept of paradigm and characterizing the classical paradigm of ecology. The classical paradigm had definite implications for conservation, and these are shown to be problematical. Therefore, we outline the contemporary paradigm in ecology and show how the science of ecology has been affected by the shift in paradigm.

Evaluating Tidal Marsh Sustainability in the Face of Sea-Level Rise: A Hybrid Modeling Approach Applied to San Francisco Bay

Background Tidal marshes will be threatened by increasing rates of sea-level rise (SLR) over the next century. Managers seek guidance on whether existing and restored marshes will be resilient under a range of potential future conditions, and on prioritizing marsh restoration and conservation activities. Methodology Building upon established models, we developed a hybrid approach that involves a mechanistic treatment of marsh accretion dynamics and incorporates spatial variation at a scale relevant for conservation and restoration decision-making. We applied this model to San Francisco Bay, using best-available elevation data and estimates of sediment supply and organic matter accumulation developed for 15 Bay subregions. Accretion models were run over 100 years for 70 combinations of starting elevation, mineral sediment, organic matter, and SLR assumptions. Results were applied spatially to evaluate eight Bay-wide climate change scenarios. Principal Findings Model results indicated that under a high rate of SLR (1.65 m/century), short-term restoration of diked subtidal baylands to mid marsh elevations (−0.2 m MHHW) could be achieved over the next century with sediment concentrations greater than 200 mg/L. However, suspended sediment concentrations greater than 300 mg/L would be required for 100-year mid marsh sustainability (i.e., no elevation loss). Organic matter accumulation had minimal impacts on this threshold. Bay-wide projections of marsh habitat area varied substantially, depending primarily on SLR and sediment assumptions. Across all scenarios, however, the model projected a shift in the mix of intertidal habitats, with a loss of high marsh and gains in low marsh and mudflats. Conclusions/Significance Results suggest a bleak prognosis for long-term natural tidal marsh sustainability under a high-SLR scenario. To minimize marsh loss, we recommend conserving adjacent uplands for marsh migration, redistributing dredged sediment to raise elevations, and concentrating restoration efforts in sediment-rich areas. To assist land managers, we developed a web-based decision support tool (www.prbo.org/sfbayslr).

Seed dispersal by Ceratogymna hornbills in the Dja Reserve, Cameroon

Seed dispersal is a process critical to the maintenance of tropical forests, yet little is known about the interactions of most dispersers with their communities. In the Dja Reserve, Cameroon, seed dispersal by the hornbills Cerato- gymna atrata, C. cylindricus and C. fistulator (Aves: Bucerotidae) was evaluated with respect to the taxonomic breadth of plants dispersed, location of seed deposition and effects on seed germination. Collectively, the three hornbill species consumed fruits from 59 tree and liana species, and likely provided dispersal for 56 of them. Hornbill-dispersed tree species composed 22% of the known tree flora of the site. Hornbill visit lengths, visit frequencies, and seed passage times indicated that few seeds were deposited beneath parent trees; in five hornbill/tree species pairings studied, 69-100% of the seeds ingested were deposited away from the parent trees. Germination trials showed that hornbill gut passage is gentle on seeds. Of 24 tree species tested, 23 germinated after passage by hornbills; of 17 planted with con- trols taken directly from trees, only four species showed evidence of inhibition of germination rate, while seven experienced unchanged germination rates and six experienced enhanced germination rates. Results suggested that Ceratogymna horn- bills rank among the most important seed dispersers found in Afrotropical forests, and they deserve increased conservation attention. Ceratogymna hornbills are likely to become increasingly important in forest regeneration as populations of larger mammalian seed dispersers (such as forest elephants and primates) diminish.

Modeling Tidal Marsh Distribution with Sea-Level Rise: Evaluating the Role of Vegetation, Sediment, and Upland Habitat in Marsh Resiliency

Tidal marshes maintain elevation relative to sea level through accumulation of mineral and organic matter, yet this dynamic accumulation feedback mechanism has not been modeled widely in the context of accelerated sea-level rise. Uncertainties exist about tidal marsh resiliency to accelerated sea-level rise, reduced sediment supply, reduced plant productivity under increased inundation, and limited upland habitat for marsh migration. We examined marsh resiliency under these uncertainties using the Marsh Equilibrium Model, a mechanistic, elevation-based soil cohort model, using a rich data set of plant productivity and physical properties from sites across the estuarine salinity gradient. Four tidal marshes were chosen along this gradient: two islands and two with adjacent uplands. Varying century sea-level rise (52, 100, 165, 180 cm) and suspended sediment concentrations (100%, 50%, and 25% of current concentrations), we simulated marsh accretion across vegetated elevations for 100 years, applying the results to high spatial resolution digital elevation models to quantify potential changes in marsh distributions. At low rates of sea-level rise and mid-high sediment concentrations, all marshes maintained vegetated elevations indicative of mid/high marsh habitat. With century sea-level rise at 100 and 165 cm, marshes shifted to low marsh elevations; mid/high marsh elevations were found only in former uplands. At the highest century sea-level rise and lowest sediment concentrations, the island marshes became dominated by mudflat elevations. Under the same sediment concentrations, low salinity brackish marshes containing highly productive vegetation had slower elevation loss compared to more saline sites with lower productivity. A similar trend was documented when comparing against a marsh accretion model that did not model vegetation feedbacks. Elevation predictions using the Marsh Equilibrium Model highlight the importance of including vegetation responses to sea-level rise. These results also emphasize the importance of adjacent uplands for long-term marsh survival and incorporating such areas in conservation planning efforts.

Phylogeny and Biogeography of the Arbutoideae (Ericaceae): Implications for the Madrean-Tethyan Hypothesis

Abstract Phylogenetic relationships within subfamily Arbutoideae (Ericaceae) were estimated using parsimony and maximum likelihood analyses of sequence data from the ITS region and part of the large subunit of nuclear ribosomal DNA. The data support the monophyly of Arctostaphylos, Arctous, and Comarostaphylis, but suggest that Arbutus is not monophyletic, with Mediterranean Basin species more closely related to the clade containing Arctostaphylos, Arctous, Comarostaphylis, Ornithostaphylos, and Xylococcus than to the western North American species of Arbutus. Calibration of branch lengths with the fossil record suggests that a vicariance event occurred among members of the Arbutoideae between western North America and the Mediterranean Basin at the Paleogene/Neogene boundary, consistent with the Madrean-Tethyan hypothesis. Communicating Editor: Kathleen A. Kron

Species-mediated soil moisture availability and patchy establishment of Pseudotsuga menziesii in chaparral

Abstract The occurrence of mature individuals of Pseudotsuga menziesii in stands of Arctostaphylos species mark the initial stages of mixed evergreen forest invasion into chaparral in central coastal California. We planted two cohorts of P. menziesii seeds at three sites under stands of two Arctostaphylos species and Adenostoma fasciculatum in order to determine whether first-year seedling emergence and survival, particularly during the regular summer drought, underlie the spatial distribution of mature trees observed in chaparral. Regardless of the chaparral species they were planted under, P. menziesii seeds that were not protected from vertebrate predation displayed very little emergence and no survival. In contrast, emergence of P. menziesii that were protected from vertebrate predators was much higher but still did not significantly differ among the three chaparral species. However, survival of protected seedlings under Arctostaphylos glandulosa was much greater than under A. fasciculatum, with intermediate survival under Arctostaphylos montana. While mortality of protected seedlings due to insect herbivory, fungal infection, and disturbance displayed no consistent patterns, summer drought mortality appeared to drive the patterns of survival of P. menziesii under the different chaparral species. These emergence, mortality, and survival data suggest that spatial patterns of P. menziesii recruitment in chaparral are driven by first-year summer drought seedling mortality, but only in years when seeds and seedlings are released from vertebrate predation pressure. Because the first-year drought mortality and survival patterns of P. menziesii seedlings differed strongly depending on the chaparral species, we examined the additional hypothesis that these patterns are associated with differences in the availability of soil moisture under different chaparral species. Both higher survival and lower drought mortality of P. menziesii seedlings were associated with higher soil water potential under Arctostaphylos stands during the summer drought, especially in the subsurface soil. The data suggest that Arctostaphylos stands, particularly stands of A. glandulosa, ameliorate xeric summer conditions to a degree that facilitates first-year establishment of P. menziesii and strongly influences spatial distribution of mature trees.

EMERGING ISSUES FOR THE RESTORATION OF TIDAL MARSH ECOSYSTEMS IN THE CONTEXT OF PREDICTED CLIMATE CHANGE

ABSTRACT There is currently a large regional effort to restore tidal marsh ecosystems in the San Francisco Bay-Delta Estuary involving the commitment of hundreds of millions of dollars and broad landscape-scale habitat manipulations. Although climate change has been on the horizon for many years, recent developments suggest that it must be taken seriously as a factor to be considered in future planning for marsh restoration efforts. Tidal marshes are vulnerable to changes in salinity and inundation rates, both of which will be affected by climate change. Restoration sites may be particularly vulnerable given unpredictable sediment inputs and newly established vegetation. Predicted shifts in snowmelt and altered runoff will change estuarine salinity patterns and could have large-scale impacts on marsh dominance, especially for freshwater marshes. Even relatively small salinity changes could lead to shifts in dominant species, with freshwater marshes being replaced by brackish marshes and brackish marshes converted to salt marsh communities. This will cause a reduction in overall estuarine plant diversity and productivity, with possible reverberations for the estuarine food web. Based on monitoring data from San Francisco Bay marshes, we predict that salinity will have a more immediate impact on tidal marsh vegetation than sea-level rise. However, sea-level rise poses a potentially greater long-term threat, depending on its rate, because the effects of inundation and a more persistent salinity regime could cause widespread marsh loss. If ice sheets in Antarctica and Greenland begin melting at rapid rates, inundation impacts could be catastrophic for coastal marshes. Given the magnitude of these potential changes, we urge the restoration and conservation management community to integrate these contingencies into adaptive management process and to join with the broader community in forging more flexible governance institutions that can respond effectively to large-scale uncertainties and trajectories as they unfold.

Effects of Litter Addition on Ectomycorrhizal Associates of a Lodgepole Pine (Pinus contorta) Stand in Yellowstone National Park

ABSTRACT Increasing soil nutrients through litter manipulation, pollution, or fertilization can adversely affect ectomycorrhizal (EM) communities by inhibiting fungal growth. In this study, we used molecular genetic methods to determine the effects of litter addition on the EM community of a Pinus contorta stand in Yellowstone National Park that regenerated after a stand-replacing fire. Two controls were used; in unmodified control plots nothing was added to the soil, and in perlite plots perlite, a chemically neutral substance, was added to maintain soil moisture and temperature at levels similar to those under litter. We found that (i) species richness did not change significantly following perlite addition (2.6 ± 0.3 species/core in control plots, compared with 2.3 ± 0.3 species/core in perlite plots) but decreased significantly (P < 0.05) following litter addition (1.8 ± 0.3 species/core); (ii) EM infection was not affected by the addition of perlite but increased significantly (P < 0.001) in response to litter addition, and the increase occurred only in the upper soil layer, directly adjacent to the added litter; and (iii) Suillus granulatus, Wilcoxina mikolae, and agaricoid DD were the dominant organisms in controls, but the levels of W. mikolae and agaricoid DD decreased significantly in response to both perlite and litter addition. The relative levels of S. granulatus and a fourth fungus, Cortinariaceae species 2, increased significantly (P < 0.01 and P < 0.05, respectively) following litter addition. Thus, litter addition resulted in some negative effects that may be attributable to moisture-temperature relationships rather than to the increased nutrients associated with litter. Some species respond positively to litter addition, indicating that there are differences in their physiologies. Hence, changes in the EM community induced by litter accumulation also may affect ecosystem function.

Community seed rain patterns and a comparison to adult community structure in a West African tropical forest

We examined the seed rain throughout a twelve month period in a lowlandtropical forest in Cameroon, West Africa, 1996–97. Traps (0.5m2, n = 216) were erected throughout a 25km2 area in 24 randomly placed clusters of nine trapseach. Fruits and seeds that landed in traps were collected every 7–10daysand classified by species and dispersal type. More than 32,000 seeds fromapproximately 200 species fell into the traps, an average of 297 seedsm−2 yr−1 Thirty species represent 82%of the total seed rain while an additional 175 species comprise the remaining18%. When we compared the adult community to the seed rain community within thesame plots, we found no apparent correlation between seed rain patterns andadult community structure for this year of study. Furthermore, only 49% of theadult tree community produced and dispersed seed into traps in this year. Morethan 100 species (52%) found in the seed rain represented long-distanceimmigrant seed rain. Seed rain was highly variable at several scales, bothspatially and seasonally, although seeds arrived in traps during eachcollectionperiod. Cluster analyses showed that traps within plots were seldom moresimilarto one another than traps between plots. While 82% of the tree species in thecommunity are thought to be animal dispersed, only 28% of all seeds that fellinto traps had been obviously “handled” (bitten, chewed, or passed)by animals. Tests for fruit and seed removal by predators or dispersers found5%or less removal rate from traps.