Recent Publications

Freita, R., Schrack, E., He, Q., Silliman, B.R., Furlong, E.B., Telles, A.C. and Costa, C.S. 2016. Consumer control of the establishment of marsh foundation plants in intertidal mudflats. Marine Ecology Progress Series. 547:79-89. (DOI: 10.3354/meps11624)


The establishment of foundation plants in bare mudflats is a critical process. While consumers are increasingly recognized to exert strong top-down control of plant performance in salt marshes, studies to date have focused on the effects of consumers on mature stands rather than on plants that are recolonizing after disturbance or where restoration has occurred. Furthermore, whether consumer-facilitated fungal infection differentially affects newly establishing plants in mudflats compared to mature stands remains poorly understood. In a salt marsh in southern Brazil, we examined the effects of herbivory by the crab Neohelice granulata and fungal infection on the survival and growth of Spartina alterniflora transplanted into mudflats. We additionally tested the effects of herbivory and fungi on newly established versus well-established stands of S. alterniflora. Highly intensive natural crab herbivory significantly reduced the development of S. alterniflora and increased its fungal infection by 50%. Light herbivory, removing only small areas of plant leaves, reduced the height growth and leaf production of directly affected tillers by about 14 to 18%, and both newly and well-established, clonally integrated stands of S. alterniflora allocated energy towards the formation of new tillers. While herbivory facilitated fungal infection and subsequent fungal damage in leaves, no significant effects of fungicide treatment or its interactions with crab grazing on S. alterniflora growth were detected, suggesting a saprophytic rather than a pathogenic role of fungi in this 3-species interaction. Here, we found that marsh grasses transplanted for restoration or those colonizing disturbance-generated mudflats may be facilitated by protection against consumers.

Li, S. and Pennings, S.C. 2016. Disturbance in Georgia salt marshes: variation across space and time. Ecosphere 7(10):e01487.10.1002/ecs2.21487.


We documented the frequency and effect on live biomass of five different types of disturbance over 14 years in creekbank and mid-marsh zones of eight salt marshes dominated by Spartina alterniflora in Georgia, USA. Wrack (floating debris) and creekbank slumping were the most common disturbances at the creekbank, and snails were the most common disturbance agent in the mid-marsh. Disturbance frequency varied among sites due to differences in plot elevation and landscape position. Wrack disturbance at the creekbank was positively correlated with plot elevation, and both initial slumping and terminal slumping of creekbank plots were negatively correlated with plot elevation. Wrack disturbance at the creekbank and snail disturbance in the mid-marsh were also most common at barrier island vs. interior marshes. Disturbance varied up to 14-fold among years. Wrack disturbance at the creekbank was negatively correlated with river discharge and sea level, and initial slumping of creekbank plots was also negatively correlated with sea level. The different disturbance types varied in their effects on end-of-year standing plant biomass. At the creekbank, wrack disturbance reduced biomass in affected plots by ~46%, but slumping did not affect biomass until the plot was totally lost. In the mid-marsh, slumping and wrack were not important disturbances, but snail disturbance reduced biomass in affected plots by ~70%. In addition, abiotic conditions (river discharge, maximum monthly temperature, sea level, and precipitation) strongly affected year-to-year variation in biomass. Across the entire landscape, fewer than a quarter of the plots on average were disturbed, and disturbance reduced overall standing biomass by ~18% in the creekbank zone and ~3% in the mid-marsh zone. Our results indicate that wrack has fairly strong effects on end-of-year biomass at the creekbank. Overall, however, variation in abiotic conditions among years had stronger effects on end-of-year standing biomass in both marsh zones than did disturbance.

Craft, C.B., Herbert, E., Li, F., Smith, D., Schubauer-Berigan, J.P., Widney, S., Angelini, C., Pennings, S.C., Medeiros, P.M., Byers, J. and Alber, M. 2016. Climate change and the fate of coastal wetlands. Wetland Science and Practice. 33(3):70-73.

O’Connell, J. and Alber, M. 2016. A smart classifier for extracting environmental data from digital image time-series: Applications for PhenoCam data in a tidal salt marsh. Environmental Modelling & Software. 84:134-139.


PhenoCams are part of a national network of automated digital cameras used to assess vegetation phenology transitions. Effectively analyzing PhenoCam time-series involves eliminating scenes with poor solar illumination or high cover of non-target objects such as water. We created a smart classifier to process images from the “GCESapelo” PhenoCam, which photographs a regularly-flooded salt marsh. The smart classifier, written in R, assigns pixels to target (vegetation) and non-target (water, shadows, fog and clouds) classes, allowing automated identification of optimal scenes for evaluating phenology. When compared to hand-classified validation images, the smart classifier identified scenes with optimal vegetation cover with 96% accuracy and other object classes with accuracies ranging from 86 to 100%. Accuracy for estimating object percent cover ranged from 74 to 100%. Pixel-classification with the smart classifier outperformed previous approaches (i.e. indices based on average color content within ROIs) and reduced variance in phenology index time-series. It can be readily adapted for other applications.

He, Q. and Silliman, B.R. 2016. Consumer control as a common driver of coastal vegetation worldwide. Ecological Monographs. 86(3):278-294. (DOI: 10.1002/ecm.1221)


Rapid, global, anthropogenic alteration of food webs in ecosystems necessitates a better understanding of how consumers regulate natural communities. We provide a global synthesis of consumer control of vegetation in coastal wetlands, where the domineering role of physical factors such as nutrient and salinity, rather than consumers, has been emphasized for decades. Using a data set of 1748 measures of consumer effects reported in 443 experiments/observations on all continents except Antarctica, we examine the generality of consumer control in salt marshes and mangroves globally. Our analyses show that salt marsh herbivores, including insects, snails, crabs, waterfowl, small mammals, and livestock, generally and often strongly suppress plant survival, aboveground biomass, and height, while their effects on plant density, belowground biomass, reproduction, and cover are more variable. These effects occur in forbs, grasses, and shrubs, and in both seedlings and adult plants. Herbivores additionally affect plant nutrient stoichiometry, and mediate plant interactions, though their effects on plant diversity are less consistent. Higher trophic levels also affect plants, as predators facilitate plant growth through trophic cascades that suppress grazer abundance and grazing rate. In mangroves, there are also signs of consumer control, though the relatively few studies available have often focused on mangrove propagules and seedlings rather than adults. Our analyses further reveal that the strength of consumer control is regulated by many physical factors. Nutrient, disturbance, and flooding, for example, amplify the negative effects of herbivores. Along latitudinal gradients, increased temperature enhances the negative effects of ectothermic herbivores, but has no effect on those of endothermic herbivores. Consumer control of coastal plants is also apparent across study methodologies: in field and laboratory settings, in observational studies, in consumer exclusion and addition experiments, in natural and transplanted plants, and in experiments of various durations. The role of consumer control in coastal vegetation worldwide highlights the need to better recognize and theoretically conceptualize both top-down and bottom-up forcing and their interactions in coastal wetlands. Improved understanding and conservation of coastal wetlands will only occur if we incorporate what the science has revealed: trophic feedbacks are an important and pervasive determinant of coastal plant communities.

Liu, W., Maung-Douglass, K., Strong, D.R., Pennings, S.C. and Zhang, Y. 2016. Geographical variation in vegetative growth and sexual reproduction of the invasive Spartina alterniflora in China. Journal of Ecology. 104:173-181. (DOI: 10.1111/1365-2745.12487)


  1. Phenotypic variation plays an important role in successful plant invasions. The spread of invasive species over large geographic ranges may be facilitated if plants can match their phenotype to local abiotic conditions. Spartina alterniflora, native to the United States, was introduced into China in 1979 and has spread over 19ᵒ of latitude along the eastern coast of China.
  2. We studied patterns in vegetative growth and sexual reproduction of Salterniflora at 22 sites at 11 geographic locations over a latitudinal gradient of ˜2000 km from Tanggu (39.05°N, high latitude) to Leizhou (20.90°N, low latitude) in China. We further evaluated the basis of phenotypic differences by growing plants from across the range in a common garden for 2 growing seasons.
  3. We found distinct latitudinal clines in plant height, shoot density and sexual reproduction across latitude. Some traits exhibited linear relationships with latitude; others exhibited hump-shaped relationships. We identified correlations between plant traits and abiotic conditions such as mean annual temperature, growing degree days, tidal range and soil nitrogen content. However, geographic variation in all but one trait disappeared in the common garden, indicating that variation largely due to phenotypic plasticity. Only a slight tendency for latitudinal variation in seed set persisted for 2 years in the common garden, suggesting that plants may be evolving genetic clines for this trait.
  4. Synthesis. The rapid spread of Spartina alterniflora (Salterniflora) in China has probably been facilitated by phenotypic plasticity in growth and reproductive traits. We found little evidence for the evolution of genetic clines in China, even though these exist for some traits in the native range. The considerable variation among clones, within provenances, that persisted in the common garden suggests a potential for the evolution of geographic clines in the future. Low fecundity of low-latitude Salterniflora populations in China might result in a slower spread at low latitudes, but Salterniflora is likely to continue to spread rapidly at high latitudes in China and into the Korean peninsula.

Angelini, C., Griffin, J.N., van de Koppel, J., Lamers, L.P.M., Smolders, A.J. P., Derkens-Hooijberg, M., van der Heide, T., and Silliman, B.R.  2016.  A Keystone Mutualism underpins resilience of a coastal ecosystem to drought.  Nature Communications, doi:10.1038/ncomms12473.  Published 18 August 2016.


Droughts are increasing in severity and frequency, yet the mechanisms that strengthen ecosystem resilience to this stress remain poorly understood. Here, we test whether positive interactions in the form of a mutualism between mussels and dominant cordgrass in salt marshes enhance ecosystem resistance to and recovery from drought. Surveys spanning 250 km of southeastern US coastline reveal spatially dispersed mussel mounds increased cordgrass survival during severe drought by 5- to 25-times. Surveys and mussel addition experiments indicate this positive effect of mussels on cordgrass was due to mounds enhancing water storage and reducing soil salinity stress. Observations and models then demonstrate that surviving cordgrass patches associated with mussels function as nuclei for vegetative re-growth and, despite covering only 0.1–12% of die-offs, markedly shorten marsh recovery periods. These results indicate that mutualisms, in supporting stress-resistant patches, can play a disproportionately large, keystone role in enhancing ecosystem resilience to climatic extremes.

O’Donnell, J.P. R. and J. F. Schalles. 2016. Examination of abiotic drivers and their influence on Spartina alterniflora biomass over a twenty-eight year period using Landsat 5 TM satellite imagery of the central Georgia Coast. Remote Sensing 8: 477-499


We examined the influence of abiotic drivers on inter-annual and phenological patterns of aboveground biomass for Marsh Cordgrass, Spartina alterniflora, on the Central Georgia Coast. The linkages between drivers and plant response via soil edaphic factors are captured in our graphical conceptual model. We used geospatial techniques to scale up in situ measurements of aboveground S. alterniflora biomass to landscape level estimates using 294 Landsat 5 TM scenes acquired between 1984 and 2011. For each scene we extracted data from the same 63 sampling polygons, containing 1222 pixels covering about 1.1 million m2. Using univariate and multiple regression tests, we compared Landsat derived biomass estimates for three S. alterniflora size classes against a suite of abiotic drivers. River discharge, total precipitation, minimum temperature, and mean sea level had positive relationships with and best explained biomass for all dates. Additional results, using seasonally binned data, indicated biomass was responsive to changing combinations of variables across the seasons. Our 28-year analysis revealed aboveground biomass declines of 33%, 35%, and 39% for S. alterniflora tall, medium, and short size classes, respectively. This decline correlated with drought frequency and severity trends and coincided with marsh die-backs events and increased snail herbivory in the second half of the study period.

Foust, C.M., Preite, V., Schrey, A.W., Alvarez, M., Robertson, M.H., Verhoeven, K.J. and Richards, C.L. 2016. Genetic and epigenetic differences associated with environmental gradients in replicate populations of two salt marsh perennials. Special Issue: Epigenetic Studies in Ecology and Evolution. Molecular Ecology. 25:1639-1652 (DOI: 10.1111/mec.13522).


While traits and trait plasticity are partly genetically based, investigating epigenetic mechanisms may provide more nuanced understanding of the mechanisms underlying response to environment. Using AFLP and methylation-sensitive AFLP, we tested the hypothesis that differentiation to habitats along natural salt marsh environmental gradients occurs at epigenetic, but not genetic loci in two salt marsh perennials. We detected significant genetic and epigenetic structure among populations and among subpopulations, but we found multilocus patterns of differentiation to habitat type only in epigenetic variation for both species. In addition, more epigenetic than genetic loci were correlated with habitat in both species. When we analysed genetic and epigenetic variation simultaneously with partial Mantel, we found no correlation between genetic variation and habitat and a significant correlation between epigenetic variation and habitat in Spartina alterniflora. In Borrichia frutescens, we found significant correlations between epigenetic and/or genetic variation and habitat in four of five populations when populations were analysed individually, but there was no significant correlation between genetic or epigenetic variation and habitat when analysed jointly across the five populations. These analyses suggest that epigenetic mechanisms are involved in the response to salt marsh habitats, but also that the relationships among genetic and epigenetic variation and habitat vary by species. Site-specific conditions may also cloud our ability to detect response in replicate populations with similar environmental gradients. Future studies analysing sequence data and the correlation between genetic variation and DNA methylation will be powerful to identify the contributions of genetic and epigenetic response to environmental gradients.

Turck, J. and V. Thompson. 2016 Revisiting the resilience of Late Archaic hunter-gatherers along the Georgia Coast. Journal of Anthropological Archaeology 43: 39-55.


This paper addresses fisher-hunter-gatherer settlement and subsistence variability of the Georgia Coast during the Archaic-Woodland transition, framed within ideas derived from Resilience Theory, and focusing on systemic shifts, or “collapse.” A critical examination of these shifts is needed to understand how communities experience change differentially, which in turn can lead to differential community resilience. Analysis of site file and radiocarbon date databases, as well as Bayesian modeling on a subset of dates, was performed at multiple scales and within distinct micro-environmental habitats. Results indicate that as sea levels dropped, there was continuity in Late Archaic occupation within deltaic areas of the coast, with intensive shellfishing occurring earlier (∼5000 cal. BP) and lasting longer (∼3500 cal. BP). In certain habitats, occupation may have continued into the following Early Woodland period. In non-deltaic areas, shellfishing occurs between 4500 and 3800 cal. BP, followed by subsistence changes and population movement. However, some of the post-3800 cal. BP occupations were fairly substantial. This differential experience of change between deltaic and non-deltaic areas was not accompanied by collapse: both areas had resilient communities. We conclude that inter-village relationships developed during the early Late Archaic period continued into the terminal portion, leading to resilience in the face of change.

Medeiros, P.M., Seidel, M., Dittmar, T., Whitman, W.B. and Moran, M.A. 2015. Drought-induced Variability in Dissolved Organic Matter Composition in a Marsh-dominated Estuary. Geophysical Research Letters. 42, 64466453. (DOI: 10.1002/2015GL064653)


The composition of dissolved organic matter (DOM) in an estuary characterized by extensive salt marsh vegetation was investigated at the molecular level using ultrahigh resolution mass spectrometry and stable carbon isotope analyses. Samples from multiple seasons covered different hydrological regimes, including anomalously low discharge conditions. The untargeted approach used allowed for identifying the DOM molecular signatures associated with different DOM sources in the estuary. DOM composition was strongly modulated by river discharge at monthly scales, with high river flow leading to significant increases in the terrigenous signature of the DOM throughout the estuary. During a severe/exceptional drought, estuarine DOM was imprinted with a distinct signature of marsh-derived compounds. The frequency of occurrence of anomalously low discharge conditions seems to have increased over the last decades. If predictions of anthropogenically-driven changes in hydroclimate are confirmed, they will likely be accompanied by changes in DOM composition in estuaries at multi-decadal time scales.

Guo, H.Y., Wieski, K., Lan, Z.J. and Pennings, S.C. 2014. Relative influence of deterministic processes on structuring marsh plant communities varies across an abiotic gradient. Oikos. 123(2):173-178.


Understanding the processes determining community structure is one of the major goals of ecological research. Both deterministic and stochastic processes may shape community structure. The challenge is to understand the relative influence of each type of process across different environmental conditions. We investigated the influence of deterministic and stochastic processes on plant community assembly in tidal marshes across a strong abiotic (salinity) gradient in three estuaries in Georgia, USA using probabilistic Raup-Crick community dissimilarity. Our results indicated that deterministic processes had an increasingly important influence on structuring plant communities in salt and brackish marshes, probably due to high heterogeneity of microhabitats produced by the interplay between abiotic stress and biotic interactions. In contrast, the influence of deterministic processes on plant community assembly decreased in tidal freshwater marshes, suggesting an increasingly important role of stochastic processes in plant community assembly in tidal freshwater marshes, probably due to the higher species richness, higher recruitment from seed, and lower levels of abiotic stress in these habitats. At the estuarine scale (across tidal freshwater, brackish and salt marshes in each estuary), our results suggested that deterministic processes also had a relatively important influence on shaping plant community structure. Our results illustrated that plant community structure in tidal marshes is influenced by both deterministic and stochastic processes, but that the relative influence of these two types of processes varies across estuarine landscapes.

Wieski, K. and Pennings, S. 2014. Latitudinal variation in resistance and tolerance to herbivory of a salt marsh shrub. Ecography. 37(8):763-769.


Interactions between plants and herbivores often vary on a geographic scale. Although theory about plant defenses and tolerance is predicated on temporal or spatial variation in herbivore damage, no single study has compared the pattern of herbivory, plant defenses and tolerance to herbivory of a single species across a latitudinal gradient. In 2002-2005 we surveyed replicate salt marshes along the Atlantic coast of the United States from Florida to Maine. At each field site we scored leaves of Iva frutescens for herbivore damage. In laboratory experiments we measured constitutive resistance and induced resistance in I. frutescens from high and low latitude sites along the Atlantic Coast. In another common garden experiment we studied tolerance to herbivory of I. frutescens from various sites. Theory predicts that constitutive resistance should matter more when damage is high, and induced resistance when herbivory is high but variable. In the field, average levels of herbivore damage, and spatial and temporal variation in herbivore damage were all greater at low versus high latitudes, indicating that constitutive as well as induced resistance should be stronger at low latitudes. Consistent with this prediction, constitutive resistance to herbivory was stronger at low latitudes. Induced resistance to herbivores was also stronger at low latitudes: it was deployed faster and lasted longer. Theory also predicts that tolerance to herbivory should be greater where average herbivory damage is greater; however, tolerance to herbivory in Iva did not depend on geographic origin. Our results emphasize the value of considering multiple ways in which plants respond to herbivores when examining geographic variation in plant-herbivore interactions.

Wieski, K. and Pennings, S.C. 2014. Climate Drivers of Spartina alterniflora Saltmarsh Production in Georgia, USA. Ecosystems. 17(3):473-484.


Tidal wetlands are threatened by global changes related not only to sea level rise but also to altered weather patterns. To predict consequences of these changes on coastal communities, it is necessary to understand how temporally varying abiotic conditions drive wetland production. In 2000-2011, we conducted annual surveys of Spartina alterniflora biomass in tidal marshes at nine sites in and around the Altamaha river estuary on the coast of Georgia, USA. End of the year live biomass was assessed in the creekbank and midmarsh zones to estimate annual net primary production (ANPP). River discharge was the most important driver of S. alterniflora ANPP, especially in creekbank vegetation. Increased river discharge reduces water column salinity, and this was most likely the proximate driver of increased production. In the midmarsh zone, the patterns were less distinct, although river discharge was again the best predictor, but maximum temperature had similar predictive ability. In contrast to results from terrestrial grasslands, we found no consistent evidence for a sharply delimited critical period for any climate driver in the tidal marsh, which indicates that plant growth was responsive to abiotic drivers at any time during the growing season. Results were broadly consistent across multiple sites within a geographic region. Our results differ from previous analyses of production in S. alterniflora marshes, which either identified oceanic drivers of S. alterniflora production or were unable to identify any drivers, likely because the low-latitude sites we studied were hotter and more affected by river discharge than those in previous studies.