The introduction and spread of smooth cordgrass (Spartina alterniflora) in South San Francisco Bay

Spartina alterniflora was first introduced into south San Francisco Bay in the 1970’s. Since that time it has spread to new areas within the south bay and is especially well established at four sites. The spread of this introduced species was evaluated by comparing its vegetative and reproductive characteristics to the native cordgrass, Spartina foliosa. The characters studied were intertidal distribution, phenology, aboveground and belowground biomass, growth rates, seed production, and germination rates. Spartina alterniflora has a wider intertidal distribution than S. foliosa and outproduced the native cordgrass in all aspects that were studied. These results indicate that the introduced species has a much better chance of becoming established in new areas than the native species, and once established, it spreads more rapidly vegetatively than the native species. Spartina alterniflora is likely to continue to spread to new areas in the bay and displace the native plant. In addition, this introduced species may effect sedimentation dynamics, available detritus, benthic algal production, wrack deposition and disturbance, habitat structure for native wetland animals, benthic invertebrate populations, and shorebird and wading bird foraging areas. *** DIRECT SUPPORT *** A01BY058 00013     

Distribution and abundance of macroinvertebrates on certain connecticut tidal marshes,with emphasis on dominant molluscs

The distribution of macroinvertebrates on Connecticut tidal marshes corresponds well with that reported for other marshes along the Atlantic and Gulf coasts of the United States. The greatest densities and biomass of the ribbed mussel,Geukensia demissa, were found on marshes in the central and western part of the state where both the annual production ofSpartina alterniflora and tidal range are large. *** DIRECT SUPPORT *** A01BY019 00011     

Effects of stem density upon sediment retention by salt marsh cord grass,Spartina alterniflora loisel

A laboratory experiment was conducted to determine whether retention of waterborne sand by salt marsh cordgrass, Spartina alterniflora Loisel, is directly related to the number of stems per unit area. Waves generated in a trough washed over a sloping beach planted with S. alterniflora sprigs: a range of stem densities (0–108 stems/m²) was examined in separate trials. The amount of sand accumulated after 60 waves is a positive nonlinear function of stem density. The greatest accretion coincided with the highest stem density tested. Shape of the beach profile was also strongly influenced by the number of stems per m².     

Phytoplankton biomass, cell diameter, and species composition in the low salinity zone of Northern San Francisco Bay estuary

Phytoplankton chlorophyll a concentration, biovolume, cell diameter, and species composition differed across the narrow, low salinity zone between 0.6‰ to 4‰ and may influence copepod food availability in the northern San Francisco Bay Estuary. The highest chlorophyll a concentrations (range 3.2–12.3 μg 1^?1), widest cell diameters (>5 μm diam), highest diatom densities and highest production rates of >10 μm diam cells occurred at the landward edge of the salinity zone in April during a strong spring tide and May during a strong neap tide. Near optimum predator/prey ratios, large prey estimated spherical diameters, and high chlorophyll a concentrations suggest these phytoplankton communities provided good food quantity and quality for the most abundant copepods, Eurytemora affinis, Sinocalanus doerrii, and Pseudodiaptomus forbesi. At the center of the zone, chlorophyll a concentrations, diatom densities, and production rates of >10 μm diam cells were lower and cell diameters were smaller than upstream. Downstream transport was accompanied by accumulation of phytoplankton with depth and tide; maximum biomass occurred on spring tide. The lowest chlorophyll a concentrations (1.4–3.6 μg 1^?) and consistently high densities (3,000–4,000 cells ml^?1) of <5 μm diam cells occurred at the seaward edge of the zone, where the green alga Nannochloris spp. and the bluegreen alga Synechococcus spp. were the most abundant phytoplankton. Low chlorophyll a concentrations and production rates of >10 μm diam cells, small prey estimated spherical diameters, and high predator/prey ratios suggested the seaward edge of the zone had poor phytoplankton food for copepodids and adult copepods. The seaward decrease in phytoplankton chlorophyll a concentration and cell diameter and shift in species composition in the low salinity zone were probably a function of an estuary-wide decrease in chlorophyll a concentration, cell diameter, and diatom density since the early 1980s that was enhanced in the low salinity zone by clam herbivory after 1987. *** DIRECT SUPPORT *** A01BY090 00008     

Emission of gaseous carbon dioxide from salt-marsh sediments and its relation to other carbon losses

Rates of CO₂ emission from bare salt-marsh sediments in areas of short and tall formSpartina alterniflora were measured monthly for 1 yr. Maximum emission rates, as high as 325 ml CO₂m^?2h^?1, were observed during summer months, while minimum rates, 10.2 ml CO₂ m^?2h^?1, were observed during the winter. An exponential function of inverse soil temperature explained most of the seasonal variability, but other factors are involved in regulating CO₂ emissions as demonstrated by rates that were higher in spring than in late summer at equivalent temperatures. Annual CO₂ emissions from bare sediments were 27.3 and 18.6 mol C m^?2 yr^?1 in communities of short and tallS. alterniflora, respectively. It was estimated that losses of dissolved inorganic carbon from the turnover of pore water, up to 14.6 mol C m^?2 yr^?1 at the creek bank (tall,S. alterniflora) site, and diffusion of CO₂ from the root system ofS. alterniflora through the culms, 12.3 to 16.2 mol C m^?2 yr^?1, could also be important pathways of carbon loss from marsh sediments. If the internal flux of CO₂ from the root system through the culm is refixed within the leaves, then the observed rate of 9.8 μI CO₂ min^?1 cm^?2 of culm cross sectional area appears to make a small but significant contribution to total photosynthesis.     

The potential impact of herbivores on the susceptibility of the marsh plantSagittaria lancifolia to saltwater intrusion in coastal wetlands

The objective of this study was to experimentally evaluate the effects of simulated herbivory on the ability of a freshwater marsh plant to recover from temporary saltwater intrusion such as can be caused by tropical storms. Sods containingSagittaria lancifolia, a dominant plant in interior coastal marshes, were manipulated in the field so as to subject plants to a pulse of 15‰ salt water for a duration of 1 wk. In addition to the exposure to salt water, some plants were also subjected to both short-term and long-term flooding treatments of 20 cm, and to simulated herbivory (clipping). Following exposure to salt water, plants were allowed to recover over the winter and were harvested the next June. Neither simulated herbivory, nor salinity, nor flooding caused any long-term effect either singly or in pairwise combinations. However, when plants were subjected to herbivory, salt water, and flooding simultaneously, reduced growth and plant death occurred. These results suggest that high levels of grazing by herbivores may increase the susceptibility of coastal marsh plants to damage from saltwater intrusion. *** DIRECT SUPPORT *** A01BY073 00002     

The grazing effects of grass shrimp,Palaemonetes pugio, on epiphytic microalgae associated withSpartina alterniflora

Experiments were performed seasonally to estimate grass shrimp,Palaemonetes pugio, grazing on the epiphytic microalgae of cordgrass,Spartina alterniflora, and to determine if grass shrimp have the potential to regulate epiphyte abundance. Grass shrimp were given access to live culms with low and high epiphytic abundance and standing dead culms collected from the streamside levee of a Louisiana salt marsh. Plexiglas frames were used to hold culms upright in aquaria and to restrict grass shrimp access to one half of each culm. We compared epiphyte biomass on the sides of culms exposed to shrimp with the corresponding unexposed sides. Epiphytes were removed from the lowest 10 cm of culms on days 0, 3, and 10, and chlorophylla (chla) measurements on each culm half were made by fluorometry. Chla biomass on culm halves not exposed to grass shrimp significantly increased over time. Percent reductions in chla on culm halves exposed to grass shrimp (calculated by subtraction from the corresponding half not exposed to shrimp) significantly increased over time for at least one culm type in all seasons. Grass shrimp caused an average 30% reduction of epiphyte biomass over 3 d and a 40% reduction over 10 d, suggesting that grass shrimp have the capability of consuming a substantial proportion of the daily production of epiphytes. Epiphytes from standing-dead culms may be more important than those from live culms at the marsh edge to the diet of grass shrimp because chla biomass was, on average, high, and standing-dead culms were seasonally abundant. Diagnostic photosynthetic pigments from selected culms, grass shrimp gut contents, and fecal pellets were identified by high performance liquid chromatography and were used to quantify the taxonomic groups of epiphytic microalgae. Results suggested that diatoms, brown algae, green algae, red algae, and cyanobacteria were present on all culms. Similarities in the pigment content of grass shrimp gut contents and fecal pellets suggested that all algal groups were ingested. Pigment data analysis could not detect a change in the composition of the microalgal assemblage associated with grass shrimp grazing. Assuming that the reduction in chla was due exclusively to grazing, grass shrimp consumed an average of 0.5–1.5 g epiphyte carbon shrimp^?1 d^?1, suggesting that grass shrimp benefit significantly from the consumption ofS. alterniflora epiphytic algae.     

Relationships of soil,plant, and microbial characteristics in silt-clay and sand,tall-formSpartina alterniflora marshes

The relationships between soil texture, plant growth, and anaerobic microbial activity in two tall-formSpartina alterniflora marshes on Sapelo Island, Georgia, were compared. The soil of one marsh was composed of typical silt-clay-sized particles; the soil of the other marsh consisted of >90% sand-sized particles. The two soils supported similar biomasses ofS. alterniflora, however, plants were taller and more robust in the silt-clay-soil than in the sand soil. Total microbial adenosine triphosphate concentrations in the silt-clay and sand soils averaged 5.71 and 1.64 μg per cm³, respectively. Seawater slurries of both soils exhibited potential for microbial sulfate reduction, methanogenesis, and glucose fermentation; rates for the processes averaged 2.03 and 0.33 nmol S-cm³ per h; 1.20 and 0.87 μmol CH₄ per cm³ per h; and 0.04 and 0.12 per min (rate constant) for the sand and silt-clay soils, respectively.     

Vesicular-arbuscular mycorrhizae in salt marshes in North Carolina

The primary objective of this research was to determine if vesicular-arbuscular (VA) mycorrhizal fungi are associated with the roots of common plant species found in North Carolina salt marshes. Root samples of Spartina alterniflora, S. patents, S. cynosuroides, Distichlis spicata, and Juncus roemerianus were collected from eight salt marsh sites. With the exception of S. alterniflora, all plant species were mycorrhizal. A greenhouse experiment was conducted to determine whether unfavorable soil conditions or inherent resistance by the plant inhibited development of mycorrhizal infection in field-collected S. alterniflora. Spartina alterniflora and S. patens were grown from seeds in soil collected from a pure stand of S. alterniflora (soil A) or a mixed stand of S. patens and D. spicata (soil P). Seedlings were harvested weekly for 8 wk, and roots were evaluated for infection by mycorrhizal fungi. Seedlings of S. patens were infected when grown for 2 wk in either soil A or soil P, indicating that soil collected from stands of S. alterniflora did not inhibit mycorrhizal infection in a susceptible host. Percent root length infected in S. patens was always greater in soil P than in soil A. Seedlings of S. alterniflora were not infected by mycorrhizal fungi in either soil A or soil P. Results of the greenhouse study indicate that S. alterniflora may be resistant to infection by vesicular-arbuscular mycorrhizal fungi.     

The Influence of Salinity on the Uptake, Distribution, and Excretion of Metals by the Smooth Cordgrass, Spartina alterniflora (Loisel.), Grown in Sediment Contaminated by Multiple Metals

The effect of salinity on the concentration of metals in Spartina alterniflora tissues and excreted salts and the distribution of metals between these compartments was tested in a greenhouse experiment. S. alterniflora stems were planted in pots of dredge sediment that was amended with a suite of six metals (Cd, Cr, Cu, Ni, Pb, and Zn). The overlying water of individual pots was maintained at treatment salinities of 0, 15, or 30 practical salinity units. The proportion of total metal distributed to the tissues and excreted salt of S. alterniflora was not significantly influenced by salinity for any of the metals. The concentration of Pb in root and leaf tissues and in excreted salt was influenced by salinity, as was the concentration of Zn in root tissue and excreted salt. Contrary to expectations, the concentration of Pb and Zn in excreted salts decreased as salinity increased. Although the influence of salinity on the concentration of Pb and Zn in S. alterniflora tissue has not been previously reported, the influence of salinity on the tissue concentrations of Pb agrees with previous research on dicots.     

Do Spur-Throated Grasshoppers, <Emphasis Type="Italic">Melanoplus</Emphasis> spp. (Orthoptera: Acrididae), Exert Top-Down Control on Smooth Cordgrass <Emphasis Type="Italic">Spartina alterniflora</Emphasis> in Northern New England?

Recently, strong top-down (consumer) control of cordgrass (Spartina alterniflora) has been demonstrated. Here, we manipulated the densities of cordgrass consumers, acridid grasshoppers (Melanoplus bivittatus and Melanoplus femurrubrum), to examine their impact on cordgrass in the Plum Island Estuary (PIE), MA, USA. After 1 month, there was no detectable effect of grasshopper density on S. alterniflora biomass and grasshoppers at the highest densities (34 individuals per square meter) consumed only ~14% of the standing stock biomass. However, significant impacts of grasshopper density on grazing damage were seen. For example, plant damage and scarring length increased by 160% and 6,156%, respectively, at the highest grasshopper densities relative to exclusion (zero grasshoppers) densities. Plant height was significantly reduced with increasing grasshopper densities, although this may be a function of leaf tip removal instead of reduced plant growth. No other strong consumers of cordgrass (e.g., Littoraria irrorata, Prokelisia marginata) were observed in PIE and we suggest that consumer regulation of cordgrass is weak in this system.     

Short- and Long-Term Vegetative Propagation of Two <Emphasis Type="Italic">Spartina</Emphasis> Species on a Salt Marsh in Southern Brazil

Spartina alterniflora and Spartina densiflora are native salt marsh plants from the Atlantic coast; their habitats in Patos Lagoon estuary (southern Brazil) are characterized by a microtidal regime (<0.5 m) and, during El Niño events, high estuarine water levels and prolonged flooding due to elevated freshwater discharge from a 200,000-km² watershed. During and between El Niño events, the vegetative propagation of these two Spartina species in the largest estuary of southern Brazil (Patos Lagoon) was evaluated by monitoring transplanted plants for 10 years (short-term study) and interpreting aerial photos of natural stands for 56 years (long-term study). During the short-term study, S. alterniflora quickly occupied mud flats (up to 208 cm year^?1) by elongation of rhizomes, whereas S. densiflora showed a modest lateral spread (up to 13 cm year^?1) and generated dense circular-shaped stands. However, moderate and strong El Niño events can promote excessive flooding and positive anomalies in the estuarine water level that reduce the lateral spread and competitive ability of S. densiflora. During the long-term study, natural stands of S. alterniflora and S. densiflora had steady lateral spread rates of 152 and 5.2 cm year^?1, respectively, over mud flats. In the microtidal marshes of the southwest Atlantic, the continuous long-term lateral expansion of both Spartina species embodies periods of intense flooding stress (moderate and strong El Niños), when there is a decrease of vegetative propagation and less stressful low water periods of fast spread over mud flats (non-El Niño periods and weak intensity El Niños).     

Biomass and productivity of three phytoplankton size classes in San Francisco Bay

Primary productivity of three size classes of phytoplankton (<5 μm, 5–22 μm, >22 μm) was measured monthly at six sites within San Francisco Bay throughout 1980. These sites in the three principal embayments were chosen to represent a range of environments, phytoplankton communities, and seasonal cycles in the estuary. Temporal variations in productivity for each size class generaly followed the seasonality of the corresponding fraction of phytoplankton biomass. The 5–22 μm size class accounted for 40 to 50% of the annual production in each embayment, but production by phytoplankton >22 μm ranged from 26% in the southern reach to 54% of total phytoplankton production in the landward embayment of the northern reach. A productivity index is derived that predicts daily productivity for each size class as a function of ambient irradiance and integrated chlorophylla in the photic zone. For the whole phytoplankton community and for each size class, this index was constant and estimated as ?0.76 g C m^?2 (g chlorophylla Einstein)^?1. The annual means of maximum carbon assimilation numbers were usually similar for the three size classes. Spatial and temporal variations in size-fractionated productivity are shown to be primarily due to differences in biomass rather than size-dependent carbon assimilation rates. *** DIRECT SUPPORT *** A01BY034 00005     

Organic matter fluxes and marsh stability in a rapidly submerging estuarine marsh

We studied organic matter cycling in two Gulf Coast tidal, nonsaline marsh sites where subsidence causes marine intrusion and rapid submergence, which mimics increased sea-level rise. The sites experienced equally rapid submergence but different degrees of marine intrusion. Vegetation was hummocked and much of the marsh lacked rooted vegetation. Aboveground standing crop and production, as measured by sequential harvesting, were low relative to other Gulf CoastSpartina patens marshes. Soil bulk density was lower than reported for healthyS. alterniflora growth but that may be unimportant at the current, moderate sulfate levels. Belowground production, as measured by sequential harvesting, was extremely fast within hummocks, but much of the marsh received little or no belowground inputs. Aboveground production was slower at the more saline site (681 g m^?2 yr^?1) than at the less saline site (1,252 g m^?2 yr^?1). Belowground production over the entire marsh surface averaged 1,401 g m^?2 yr^?1 at the less saline site and 585 g m^?2 yr^?1 at the more saline site. Respiration, as measured by CO₂ emissions in the field and corrected for CH₄ emissions, was slower at the less saline site (956 g m^?2 yr^?1) than at the more saline site (1,438 g m^?2 yr^?1), reflecting greater contributions byS. alterniflora at the more saline site which is known to decompose more rapidly thanS. patens. Burial of organic matter was faster at the less saline site (796 g m^?2 yr^?1) than at the more saline site (434 g m^?2, yr^?1), likely in response to faster production and slower decomposition at the less saline site. Thus vertical accretion was faster at the less saline site (1.3 cm yr^?1) than at the more saline site (0.85 cm yr^?1); slower vertical accretion increased flooding at the more saline site. More organic matter was available for export at the less saline site (1,377 g m^?2 yr^?1) than at the more saline site (98 g m^?2 yr^?1). These data indicated that organic matter production decreased and burial increased in response to greenhouse-like conditions brought on by subsidence. *** DIRECT SUPPORT *** A01BY069 00016     

Aboveground Biomass Patterns of Dominant Spartina Species and Their Relationship with Selected Abiotic Variables in Argentinean SW Atlantic Marshes

Salt marsh zonation patterns generate different abiotic and biotic conditions that can accentuate species inherent differences in primary production and biomass. In South West Atlantic marshes, there are two Spartina species: Spartina alterniflora in the low intertidal and Spartina densiflora in the high intertidal. These two species are generally found in all marshes but with different dominance: In some marshes, the S. densiflora zone occupies higher extents, and in others, the S. alterniflora zone is the one that prevails. We found through field sampling that, in six studied marshes, there is greater S. densiflora live and total (i.e., dead+live) aboveground biomass (g m^?2) in the marshes dominated by S. densiflora than in the ones dominated by S. alterniflora. Spartina alterniflora had similar aboveground biomass in the six marshes, regardless of the dominance of each species. When comparing the two Spartina species within each marsh, S. densiflora had greater live and total biomass in the marshes it dominates. In the marshes dominated by S. alterniflora, both species had similar live and total biomass. In all marshes, there was greater dead S. densiflora biomass. A multivariate analysis using selected abiotic factors (i.e., salinity, latitude, and tidal amplitude) showed that S. alterniflora aboveground biomass patterns are mainly correlated with salinity, while S. densiflora live biomass is mainly correlated with salinity and latitude, dead biomass with salinity and tidal amplitude, and total biomass with salinity alone. We conclude that in S. densiflora dominated marshes, the main processes of that species zone (i.e., nutrient accumulation) will be accentuated because of its higher biomass. We also conclude that climatic conditions, in combination with specific Spartina biotic and ambient abiotic parameters, can affect marsh ecological functions.     

Egg Deposition by Atlantic Silverside, Menidia menidia: Substrate Utilization and Comparison of Natural and Altered Shoreline Type

Egg deposition by the intertidal spawning fish Atlantic silverside (Menidia menidia) was compared among six shoreline types (Spartina alterniflora, Phragmites australis, sandy beach, riprap, riprap-sill, and bulkhead) and various substrates. In spring 2010, M. menidia egg density was measured daily near Roosevelt Inlet, Delaware Bay, USA. Over 3,000,000 eggs were collected during 50 sampling days. Eggs were deposited at all six shoreline types, with >93?% of eggs collected from S. alterniflora shorelines. Choice of substrate for egg attachment was similar across shoreline types with >91?% of eggs collected from filaments of the green alga Enteromorpha spp., a disproportionately high utilization rate in comparison with Enteromorpha spp.'s relative coverage. This study demonstrates that S. alterniflora shoreline, in association with Enteromorpha spp., is the preferred spawning habitat for M. menidia and that hardened shorelines and shorelines inhabited by P. australis support substantially reduced egg densities.     

Ecological Impacts of Macroalgal Blooms on Salt Marsh Communities

Despite excessive growth of macroalgae in estuarine systems, little research has been done to examine the impacts of increased algal biomass that drifts into nearby salt marshes and accumulates on intertidal flats. The accumulation of macroalgal mats and subsequent decomposition-related releases of limiting nutrients may potentially alter marsh communities and impact multiple trophic levels. We conducted a 2-year in situ study, as well as laboratory mesocosm experiments, to determine the fate of these nutrients and any bottom-up impacts from the blooms on the dominant salt marsh plant (Spartina alterniflora) and herbivores. Mesocosm results showed that macroalgal decomposition had a positive impact on sediment nitrogen concentrations, as well as S. alterniflora growth rates. In contrast, our in situ results suggested that S. alterniflora growth was hindered by the presence of macroalgal mats. From our results, we suggest that macroalgal accumulation and subsequent release of nitrogen during decomposition may be beneficial in nitrogen limited areas. However, as marshes are becoming increasingly eutrophic, releasing lower marsh plants from nitrogen limitation, this accumulation of macroalgal biomass may hinder S. alterniflora growth through smothering and breakage of culms. As macroalgal blooms are predicted to intensify with rising temperatures and increased eutrophication, the ecological impacts associated with these changes need to be continuously monitored in order to preserve these fragile ecosystems.     

Living on the Edge: Increasing Patch Size Enhances the Resilience and Community Development of a Restored Salt Marsh

Foundation species regulate communities by reducing environmental stress and providing habitat for other species. Successful restoration of biogenic habitats often depends on restoring foundation species at appropriate spatial scales within a suitable range of environmental conditions. An improved understanding of the relationship between restoration scale and environmental conditions has the potential to improve restoration outcomes for many biogenic habitats. Here, we identified and tested whether inundation/exposure stress and spatial scale (patch size) can interactively determine (1) survival and growth of a foundation species, Spartina alterniflora and (2) recruitment of supported fauna. We planted S. alterniflora and artificial mimics in large and small patches at elevations above and below local mean sea level (LMSL) and monitored plant survivorship and production, as well as faunal recruitment. In the first growing season, S. alterniflora plant survivorship and stem densities were greater above LMSL than below LMSL regardless of patch size, while stem height was greatest in small patches below LMSL. By the third growing season, S. alterniflora patch expansion was greater above LMSL than below LMSL, while stem densities were higher in large patches than small patches, regardless of location relative to LMSL. Unlike S. alterniflora, which was more productive above LMSL, sessile marine biota recruitment to mimic plants was higher in patches below LMSL than above LMSL. Our results highlight an ecological tradeoff at ~LMSL between foundation species restoration and faunal recruitment. Increasing patch size as inundation increases may offset this tradeoff and enhance resilience of restored marshes to sea-level rise.     

Sporormiella as a proxy for non-mammalian herbivores in island ecosystems

Spores of the dung-fungi Sporormiella are routinely used as a proxy for past megaherbivore biomass and to pinpoint the timing of extinctions. Further ecological insight can also be gained into the impacts that followed initial human arrival in a region through correlation of spore abundance with other proxies (e.g. pollen, charcoal). Currently, the use of Sporormiella as a palaeoecological proxy has been restricted to landmasses where large-herbivore guilds are dominated by mammals. Here, we use New Zealand as a case study to show that the method can also be applied effectively to islands dominated by large avian herbivores. We examine 44 dung samples from 7 localities to show that Sporormiella spores were widely distributed in the dung of endemic avian herbivores (South Island takahe (Porphyrio hochstetteri), kakapo (Strigops habroptilus), and several species of extinct moa, identified by ancient DNA analysis). In addition, we show that Sporormiella spores in a forest soil core from the Murchison Mountains, South Island, accurately trace the post-settlement decline of native avian herbivores, and combined with high-resolution radiocarbon dating reveal severely reduced local herbivore populations by the late 17th Century AD. The spores also trace the subsequent spread of Red deer (Cervus elaphus) introduced to the area in the early 20th Century AD. Our results suggest Sporormiella spores may provide a useful new tool for examining extinctions on numerous islands where terrestrial herbivore guilds were dominated by birds or reptiles. Our findings also highlight the need to consider entire herbivore communities (including birds and reptiles) when examining Late Pleistocene continental Sporormiella records, where the focus has often been tracing the decline of populations of large mammals.