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Steve Pennings, Field Director / Associate Professor

Research Emphasis:

Plant-herbivore interactions and plant community ecology. My students and I study how and why interactions between salt marsh plants and their herbivores vary on multiple spatial scales, and how these affect the marsh plant community. We are also interested in the broader web of processes (including top-down and bottom-up forces) that affect plant-herbivore interactions. We also study how competition between plant species is affected by multiple physical stressors in estuarine habitats, and how these interactions create spatial pattern in the plant community and spatial and temporal variation in primary productivity.

GCE Committees:

GCE Predator Exclusion Experiment Committee, GCE Disturbance Committee, GCE Flux Tower Committee, GCE Executive Committee, High Marsh Experiment Committee, LTER Network-wide Committee Members, SALTEx Experiment Committee, Salinity Monitoring Committee, Taxonomic Database Committee

Contact Information:

Primary Organization:  University of Houston

Mailing Address:

Dr. Steven C. Pennings
Department of Biology and Biochemistry
University of Houston
Houston, Texas 77204-5513

Office Phone:  (713) 743-2989

FAX Number:  (713) 743-2622

E-Mail:  scpennin@central.uh.edu

Web Page:  https://uh.edu/nsm/biology-biochemistry/people/profiles/steven-pennings/

Other Identity Systems:

ORCID:  orcid.org/0000-0003-4757-7125

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GCE Data Sets:

PLT-GCED-2407 (Soil temperature at GCE core monitoring sites in the winter of 2019-2020)

INV-GCEM-2409 (Fall 2023 crab population monitoring: mid-marsh and creek bank abundance based on crab hole counts at GCE marsh, monitoring sites 1-10)

INV-GCEM-2404a (Survey of adult and juvenile periwinkle snail (Littoraria irrorata) density in mid-marsh and creekbank plots at GCE LTER study sites in October 2023.)

INV-GCEM-2404 (Yearly survey of barnacle settlement near creekbank plots at GCE LTER study sites in October 2023)

PLT-GCEM-2311b (Fall 2023 plant monitoring survey -- biomass calculated from shoot height and flowering status of plants in permanent plots at GCE sampling sites 1-10)

(most recent displayed - view all)

GCE Publications and Presentations: (custom bibliography)

Journal Articles

Sharp, S., Davidson, K., Angelini, C., Fischman, H., Pennings, S.C., Fowler, M.S. and Griffin, J. 2024. Large grazers suppress a foundational plant and reduce soil carbon concentration in eastern US saltmarshes. Journal of Ecology. (DOI: 10.1111/1365-2745.14407)

Smith, C.S., Zhang, S., Hensel, M.S., Pennings, S.C. and Silliman, B.R. 2024. Long-term data reveal that grazer density mediates climatic stress in salt marshes. Ecology. (DOI: 10.1002/ecy.4323)

Smith, R.S., Pennings, S.C., Alber, M., Craft, C.B. and Byers, J. 2024. The resistance of Georgia coastal marshes to hurricanes. Ecosphere. 15(4). (DOI: 10.1002/ecs2.4821)

Thompson, M., Pennings, S.C., Schubauer-Berigan, J.P., Herbert, E., Costomiris, G. and Craft, C.B. 2024. Resistance and resilience: Tidal freshwater marsh response and recovery to acute and chronic saltwater intrusion. Estuarine, Coastal and Shelf Science. (DOI: 10.1016/j.ecss.2024.108911)

Wang, J., Guo, H., Alber, M. and Pennings, S.C. 2024. Variance reflects resilience to disturbance along a stress gradient: experimental evidence from coastal marshes. Ecology. 2024:e4241. (DOI: https://doi.org/10.1002/ecy.4241)

Adams, T., Vu, H. and Pennings, S.C. 2022. Variation in Densities of the Salt Marsh Katydid Orchelimum fidicinium over Space and Time. Estuaries and Coasts. 45:260-271. (DOI: https://doi.org/10.1007/s12237-021-00953-y)

Li, F., Angelini, C., Byers, J., Craft, C.B. and Pennings, S.C. 2022. Responses of a tidal freshwater marsh plant community to chronic and pulsed saline intrusion. Journal of Ecology. 110:1508-1524. (DOI: 10.1111/1365-2745.13885)

Nie, M., Liu, W., Pennings, S.C. and Li, B. 2022. Lessons from the invasion of Spartina alterniflora in coastal China. Ecology. 104:e3874. (DOI: 10.1002/ecy.3874)

Simon, J., Hopkinson, B.M. and Pennings, S.C. 2022. Insights into Salt Marsh Plant Community Distributions Through Computer Vision and Structural Equation Modeling. Estuaries and Coasts. 46:431-449. (DOI: 10.1007/s12237-022-01147-w)

Chen, X., Liu, W., Pennings, S.C. and Zhang, Y. 2021. Plasticity and selection drive hump-shaped latitudinal patterns of flowering phenology in an invasive intertidal plant. Ecology. 102. (DOI: 10.1002/ecy.3311)

Liu, W. and Pennings, S.C. 2021. Variation in synchrony of production among species, sites and intertidal zones in coastal marshes. Ecology. (DOI: 10.1002/ECY.3278)

Parashar, J., Bhandarkar, S.M., Simon, J., Hopkinson, B. and Pennings, S.C. 2021. Estimation of Abundance and Distribution of Salt Marsh Plants from Images Using Deep Learning. Proceedings of the 25th International Conference on Pattern Recognition (ICPR). (DOI: 10.1109/ICPR48806.2021.9412264)

Seer, F., Putze, G., Pennings, S.C. and Zimmer, M. 2021. Drivers of litter mass loss and faunal composition of detritus patches change over time. Ecology and Evolution. 11:9642-9651. (DOI: 10.1002/ece3.7787)

Vu, H. and Pennings, S.C. 2021. Directional movement of consumer fronts associated with creek heads in salt marshes. Ecology. 102(9):e03447. (DOI: 10.1002/ecy.3447)

Wu, F., Pennings, S.C., Ortals, C., Ruiz, J., Farrell, W.R., McNichol, S.M., Angelini, C., Spivak, A.C., Alber, M. and Tong, C. 2021. Disturbance is complicated: headward-eroding saltmarsh creeks produce multiple responses and recovery trajectories. Limnology & Oceanography. 67:S86-S100. (DOI: 10.1002/lno.11867)

Zhang, Y., Pennings, S.C., Liu, Z., Li, B. and Wu, J. 2021. Consistent pattern of higher lability of leaves from high latitudes for both native Phragmites australis and exotic Spartina alterniflora. Functional Ecology. 35:2084-2093. (DOI: https://doi. org/10.1111/1365-2435.13826)

Zinnert, J.C., Nippert, J.B., Rudgers, J.A., Pennings, S.C., Gonzalez, G., Alber, M., Baer, S.G., Blair, J.M., Burd, A.B., Collins, S.L., Craft, C.B., Di Iorio, D., Dodds, W.K., Groffman, P.M., Herbert, E., Hladik, C.M., Li, F., Litvak, M., Newsome, S., O'Donnell, J., Pockman, W.T., Schalles, J.F. and Young, D.R. 2021. State Changes: Insights from the U.S. Long Term Ecological Research Network. Ecosphere. (DOI: 10.1002/ecs2.3433)

Liu, W., Chen, X., Strong, D.R., Pennings, S.C., Kirwan, M.L., Chen, X. and Zhang, Y. 2020. Climate and geographic adaptation drive latitudinal clines in biomass of a widespread saltmarsh plant in its native and introduced ranges. Limnology & Oceanography. 65:1399-1409. (DOI: 10.1002/lno.11395)

Liu, W., Zhang, Y., Chen, X., Maung-Douglass, K., Strong, D.R. and Pennings, S.C. 2020. Contrasting plant adaptation strategies to latitude in the native and invasive range of Spartina alterniflora. New Phytologist. 226:623-634. (DOI: 10.1111/nph.16371)

Nahrawi, H.B., Leclerc, M.Y., Pennings, S.C., Zhang, G., Singh, N. and Pahari, R. 2020. Impact of tidal inundation on the net ecosystem exchange in daytime conditions in a salt marsh. Agricultural and Forest Meteorology. 294:108133. (DOI: https://doi.org/10.1016/j.agrformet.2020.108133)

O'Connell, J.L., Alber, M. and Pennings, S.C. 2020. Microspatial differences in soil temperature cause phenology change on par with long-term climate warming in salt marshes. Ecosystems. 23:498–510. (DOI: https://doi.org/10.1007/s10021-019-00418-1)

Zhang, Y., Li, B., Wu, J. and Pennings, S.C. 2020. Contrasting latitudinal clines of nematode diversity in Spartina alterniflora salt marshes between native and introduced ranges. Diversity and Distributions. 26:623-631. (DOI: 10.1111/ddi.13054)

Li, F. and Pennings, S.C. 2019. Response and Recovery of Low-Salinity Marsh Plant Communities to Presses and Pulses of Elevated Salinity. Estuaries and Coasts. 42:708-718. (DOI: 10.1007/s12237-018-00490-1)

Li, S., Xie, T., Pennings, S.C., Wang, Y., Craft, C.B. and Hu, M. 2019. A comparison of coastal habitat restoration projects in China and the United States. Scientific Reports. 9(14388). (DOI: 10.1038/s41598-019-50930-6)

Liu, W. and Pennings, S.C. 2019. Self-thinning and size-dependent flowering of the grass Spartina alterniflora across space and time. Functional Ecology. 33:1830-1841. (DOI: 10.1111/1365-2435.13384)

Widney, S., Smith, D., Herbert, E., Schubauer-Berigan, J.P., Li, F., Pennings, S.C. and Craft, C.B. 2019. Chronic but not acute saltwater intrusion leads to large release of inorganic N in a tidal freshwater marsh. Science of the Total Environment. 695. (DOI: https://doi.org/10.1016/j.scitotenv.2019.133779)

Zhang, C., Mishra, D. and Pennings, S.C. 2019. Mapping salt marsh soil properties using imaging spectroscopy. ISPRS Journal of Photogrammetry and Remote Sensing. 148:221-234. (DOI: 10.1016/j.isprsjprs.2019.01.006)

Zhang, Y., Pennings, S.C., Li, B. and Wu, J. 2019. Biotic homogenization of wetland nematode communities by exotic Spartina alterniflora in China. Ecology. 100. (DOI: 10.1002/ecy.2596)

Gehman, A., McLenaghan, N.A., Byers, J., Alexander, C.R. Jr., Pennings, S.C. and Alber, M. 2018. Effects of small-scale armoring and residential development on the salt marsh-upland ecotone. Estuaries and Coasts. 41(1):54-67. (DOI: 10.1007/s12237-017-0300-8)

Li, F. and Pennings, S.C. 2018. Responses of tidal freshwater and brackish marsh macrophytes to pulses of saline water simulating sea level rise and reduced discharge. Wetlands. 38:885-891. (DOI: 10.1007/s13157-018-1037-2)

Li, S., Hopkinson, C.S., Schubauer-Berigan, J.P. and Pennings, S.C. 2018. Climate drivers of Zizaniopsis miliacea biomass in a Georgia, U.S.A. tidal fresh marsh. Limnology and Oceanography. 63:2266-2276. (DOI: 10.1002/lno.10937)

Lin, W.-T. and Pennings, S.C. 2018. Predator-prey interactions in a ladybeetle-aphid system depend on spatial scale. Ecology and Evolution. 8:6537-6546. (DOI: 10.1002/ece3.4117)

Peng, D., Chen, L., Pennings, S.C. and Zhang, Y. 2018. Using a marsh organ to predict future plant communities in a Chinese estuary invaded by an exotic grass and mangrove. Limnology and Oceanography. 63:2595-2605. (DOI: 10.1002/lno.10962)

Vu, H. and Pennings, S.C. 2018. Predators mediate above- vs belowground herbivory in a salt marsh crab. Ecosphere. 9(2):e02107. (DOI: 10.1002/ecs2.2107)

Li, S. and Pennings, S.C. 2017. Timing of disturbance affects biomass and flowering of a saltmarsh plant and attack by stem-boring herbivores. Ecosphere. 8(2):9. (DOI: 10.1002/ecs2.1675)

Liu, W., Strong, D.R., Pennings, S.C. and Zhang, Y. 2017. Provenance-by-environment interaction of reproductive traits in the invasion of Spartina alterniflora in China. Ecology. 98(6):1591-1599. (DOI: 10.1002/ecy.1815)

Vu, H., Wieski, K. and Pennings, S.C. 2017. Ecosystem engineers drive creek formation in salt marshes. Ecology. 98(1):162-174.

Zengel, S., Weaver, J., Pennings, S.C., Silliman, B.R., Deis, D., Montague, C.L., Zimmerman, A., Rutherford, N. and Nixon, Z. 2017. Five years of Deepwater Horizon oil spill effects on marsh periwinkles Littoraria irrorata. Marine Ecology Progress Series. 576:135-144. (DOI: 10.3354/meps11827)

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.

Li, S. and Pennings, S.C. 2016. Disturbance in Georgia salt marshes: variation across space and time. Ecosphere. 7(10):e01487. (DOI: 10.1002/ecs2.1487)

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)

Guo, H., Chamberlain, S.A., Elhaik, E., Jalli, I., Lynes, A.R., Marczak, L., Sabath, N., Vargas, A., Wieski, K., Zelig, E. and Pennings, S.C. 2015. Geographic Variation in Plant Community Structure of Salt Marshes: Species, Functional and Phylogenetic Perspectives. PLOS ONE. 10(5). (DOI: 10.1371/journal.pone.0127781)

Hübner, L., Pennings, S.C. and Zimmer, M. 2015. Sex- and habitat-specific movement of an omnivorous semi-terrestrial crab controls habitat connectivity and subsidies: A multi-parameter approach. Oecologia. 3/2015:17. (DOI: 10.1007/s00442-015-3271-0)

Guo, H., Wieski, K., Lan, Z. 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. (DOI: 10.1111/j.1600-0706.2013.00425.x)

He, Q., Bertness, M.D., Bruno, J.F., Li, B., Chen, G., Coverdale, T.C., Alteiri, A., Bai, J., Sun, T., Pennings, S.C., Liu, J., Ehrlich, P.R. and Cui, B. 2014. Economic development and coastal ecosystem change in China. Scientific Reports. 4(5995):9. (DOI: 10.1038/srep05995)

Pennings, S.C., McCall, B.D. and Hooper-Bui, L. 2014. Effects of oil spills on terrestrial arthropods in coastal wetlands. Bioscience. 64:789-795. (DOI: 10.1093/biosci/biu118)

Wieski, K. and Pennings, S.C. 2014. Climate Drivers of Spartina alterniflora Saltmarsh Production in Georgia, USA. Ecosystems. 17(3):473-484. (DOI: 10.1007/s10021-013-9732-6)

Wieski, K. and Pennings, S.C. 2014. Latitudinal variation in resistance and tolerance to herbivory of a salt marsh shrub. Ecography. 37:763-769. (DOI: 10.1111/ecog.00498)

Guo, H., Zhang, Y., Lan, Z. and Pennings, S.C. 2013. Biotic interactions mediate the expansion of black mangrove (Avicennia germinans) into salt marshes under climate change. Global Change Biology. 19:2765 - 2744. (DOI: 10.1111/gcb.12221)

Ho, C.-K. and Pennings, S.C. 2013. Preference and performance in plant-herbivore interactions across latitude – a study in U.S. Atlantic salt marshes. PLoS One. 8(3). (DOI: 10.1371/journal.pone.0059829)

Marczak, L., Wieski, K., Denno, R. and Pennings, S.C. 2013. Importance of local vs. geographic variation in salt marsh plant quality for arthropod herbivore communities. Journal of Ecology. 101:1169-1182. (DOI: 10.1111/1365-2745.12137)

Schalles, J.F., Hladik, C.M., Lynes, A.R. and Pennings, S.C. 2013. Landscape estimates of habitat types, plant biomass, and invertebrate densities in a Georgia salt marsh. Special Issue: Coastal Long Term Ecological Research. Oceanography. 26:88-97. (DOI: 10.5670/oceanog.2013.50)

Treplin, M., Pennings, S.C. and Zimmer, M. 2013. Decomposition of leaf litter in a U.S. saltmarsh is driven by dominant species, not species complementarity. Wetlands. 33:83-89. (DOI: 10.1007/s13157-012-0353-1)

Ewers, C., Beiersdorf, A., Wieski, K., Pennings, S.C. and Zimmer, M. 2012. Predator/prey-interactions promote decomposition of low-quality detritus. Wetlands. 32:931-938. (DOI: 10.1007/s13157-012-0326-4)

Gough, L., Gross, K.L., Cleland, E.E., Clark, C.M., Collins, S.L., Fargione, J.E., Pennings, S.C. and Suding, K.N. 2012. Incorporating clonal growth form clarifies the role of plant height in response to nitrogen addition. Oecologia. 169:1053-1062. (DOI: 10.1007/s00442-012-2264-5)

Guo, H. and Pennings, S.C. 2012. Mechanisms mediating plant distributions across estuarine landscapes in a low-latitude tidal estuary. Ecology. 93(1):90-100. (DOI: 10.1890/11-0487.1)

Guo, H. and Pennings, S.C. 2012. Post-mortem ecosystem engineering by oysters creates habitat for a rare marsh plant. Oecologia. 170:789-798. (DOI: 10.1007/s00442-012-2356-2)

Jimenez, J.M., Wieski, K., Marczak, L., Ho, C.-K. and Pennings, S.C. 2012. Effects of an omnivorous katydid, salinity, and nutrients on a planthopper-Spartina food web. Estuaries and Coasts. 35:475-485. (DOI: 10.1007/s12237-011-9458-7)

McCall, B.D. and Pennings, S.C. 2012. Disturbance and recovery of salt marsh arthropod communities following BP Deepwater Horizon oil spill. PLoS ONE. (DOI: 10.1371/journal.pone.0032735)

McCall, B.D. and Pennings, S.C. 2012. Geographic variation in salt marsh structure and function. Oecologia. 170:777-787. (DOI: 10.1007/s00442-012-2352-6)

Cleland, E.E., Clark, C.M., Collins, S.L., Fargione, J.E., Gough, L., Gross, K.L., Pennings, S.C. and Suding, K.N. 2011. Patterns of trait convergence and divergence among native and exotic species in herbaceous plant communities are not modified by nitrogen enrichment. Journal of Ecology. 99:1327-1338. (DOI: 10.1111/j.1365-2745.2011.01860.x)

Marczak, L., Ho, C.-K., Vu, H., Denno, R. and Pennings, S.C. 2011. Latitudinal variation in top-down and bottom-up control of a salt marsh food web. Ecology. 92:276-281. (DOI: 10.1890/10-0760.1)

Ho, C.-K., Pennings, S.C. and Carefoot, T.H. 2010. Is Diet Quality an Overlooked Mechanism for Bergmann's Rule? The American Naturalist. 175(2):269-276. (DOI: 10.1086/649583)

Robinson, J.D., Diaz-Ferguson, E., Poelchau, M., Pennings, S.C., Bishop, T.D. and Wares, J.P. 2010. Multiscale Diversity in the Marshes of the Georgia Coastal Ecosystems LTER. Estuaries and Coasts. 33(4):865-877. (DOI: 10.1007/s12237-009-9188-2)

Wieski, K., Guo, H., Craft, C.B. and Pennings, S.C. 2010. Ecosystem functions of tidal fresh, brackish and salt marshes on the Georgia coast. Estuaries and Coasts. 33:161-169. (DOI: 10.1007/s12237-009-9230-4)

Craft, C.B., Clough, J., Ehman, J., Joye, S.B., Park, R., Pennings, S.C., Guo, H. and Machmuller, M. 2009. Forecasting the effects of accelerated sea level rise on tidal marsh ecosystem services. Frontiers in Ecology and the Environment. 7(2):73-78. (DOI: 10.1890/070219)

Craft, C.B., Clough, J., Ehman, J., Joye, S.B., Park, R., Pennings, S.C., Guo, H. and Machmuller, M. 2009. SLR and ecosystem services: a response to Kirwan and Guntenspergen. Frontiers in Ecology and the Environment. 7:127-128.

Pennings, S.C., Ho, C.-K., Salgado, C.S., Wieski, K., Davé, N.B., Kunza Vargas, A.E. and Wason, E.L. 2009. Latitudinal variation in herbivore pressure in Atlantic Coast salt marshes. Ecology. 90(1):183-195. (DOI: 10.1890/08-0222.1)

Cleland, E.E., Clark, C.M., Collins, S.L., Fargione, J.E., Gough, L., Gross, K.L., Milchunas, D.G., Pennings, S.C., Bowman, W.D., Burke, I.C., Lauenroth, G.P., Robertson, J.C., Simpson, J.C., Tilman, D. and Suding, K.N. 2008. Species responses to nitrogen fertilization in herbaceous plant communities, and associated species traits. Ecology. 89:1175.

Collins, S.L., Suding, K.N., Cleland, E.E., Batty, M., Pennings, S.C., Gross, K.L., Grace, J.B., Gough, L., Fargione, J.E. and Clark, C.M. 2008. Rank clocks and plant community dynamics. Ecology. 89(12):3534-3541. (DOI: 10.1890/07-1646.1)

Ho, C.-K. and Pennings, S.C. 2008. Consequences of omnivory for trophic interactions on a salt-marsh shrub. Ecology. 89(6):1714-1722. (DOI: 10.1890/07-1069.1)

Kunza, A.E. and Pennings, S.C. 2008. Patterns of plant diversity in Georgia and Texas salt marshes. Estuaries and Coasts. 31:673-681.

McFarlin, C.R., Brewer, J.S., Buck, T.L. and Pennings, S.C. 2008. Impact of fertilization on a salt marsh food web in Georgia. Estuaries and Coasts. 31:313-325.

Pennings, S.C. and Simpson, J.C. 2008. Like herbivores, parasitic plants are limited by host nitrogen content. Plant Ecology. 196:245-250. (DOI: 10.1007/s11258-007-9348-z)

Wason, E.L. and Pennings, S.C. 2008. Grasshopper (Orthoptera: Tettigoniidae) species composition and size across latitude in Atlantic Coast salt marshes. Estuaries and Coasts. 31:335-343.

Clark, C.M., Cleland, E.E., Collins, S.L., Fargione, J.E., Gough, L., Gross, K.L., Pennings, S.C., Suding, K.N. and Grace, J.B. 2007. Environmental and plant community determinants of species loss following nitrogen enrichment. Ecology Letters. 10:596-607.

Pennings, S.C., Zimmer, M., Dias, N., Sprung, M., Davé, N., Ho, C.-K., Kunza, A., McFarlin, C., Mews, M., Pfauder, A. and Salgado, C. 2007. Latitudinal variation in plant-herbivore interactions in European salt marshes. Oikos. 116:543-549.

Pennings, S.C. and Silliman, B.R. 2005. Linking biogeography and community ecology: latitudinal variation in plant-herbivore interaction strength. Ecology. 86:2310-2319.

Pennings, S.C., Clark, C.M., Cleland, E.E., Collins, S.L., Gough, L., Gross, K.L., Milchunas, D.G. and Suding, K.N. 2005. Do individual plant species show predictable responses to nitrogen addition across multiple experiments? OIKOS. 110:547-555.

Pennings, S.C., Grant, M.-B. and Bertness, M.D. 2005. Plant zonation in low-latitude salt marshes: disentangling the roles of flooding, salinity and competition. Journal of Ecology. 93:159-167.

Richards, C.L., Pennings, S.C. and Donovan, L.A. 2005. Habitat range and phenotypic variation in salt marsh plants. Plant Ecology. 176:263-273.

Salgado, C.S. and Pennings, S.C. 2005. Latitudinal variation in palatability of salt-marsh plants: are differences constitutive? Ecology. 86(6):1571-1579.

Suding, K.N., Collins, S.L., Gough, L., Clark, C.M., Cleland, E.E., Gross, K.L., Milchunas, D.G. and Pennings, S.C. 2005. Functional- and abundance-based mechanisms explain diversity loss due to N fertilization. Proceedings of the National Academy of Sciences, U.S.A. 102:4387-4392.

Goranson, C.E., Ho, C.-K. and Pennings, S.C. 2004. Environmental gradients and herbivore feeding preferences in coastal salt marshes. Oecologia. 140:591-600.

Zimmer, M., Pennings, S.C., Buck, T.L. and Carefoot, T.H. 2004. Salt Marsh Litter and Detritivores: A Closer Look at Redundancy. Estuaries. 27(5):753–769.

Buck, T.L., Breed, G.A., Pennings, S.C., Chase, M.E., Zimmer, M. and Carefoot, T. 2003. Diet choice in an omnivorous salt marsh crab: different food types, claw allometry, and habitat complexity. Journal of Experimental Marine Biology and Ecology. 292:103-116. (DOI: 10.1016/S0022-0981(03)00146-1)

Callaway, R.M., Pennings, S.C. and Richards, C.L. 2003. Phenotypic plasticity and interactions among plants. Ecology. 84(5):1115-1128.

Pennings, S.C., Selig, E.R., Houser, L.T. and Bertness, M.D. 2003. Geographic variation in positive and negative interactions among salt marsh plants. Ecology. 84:1527-1538.

Pennings, S.C. and Callaway, R.M. 2002. Parasitic plants: parallels and contrasts with herbivores. Oecologia. 131:479-489.

Pennings, S.C., Stanton, L.E. and Brewer, J.S. 2002. Nutrient effects on the composition of salt marsh plant communities along the southern Atlantic and Gulf Coasts of the United States. Estuaries. 25:1164-1173.

Zimmer, M., Pennings, S.C., Buck, T.L. and Carefoot, T.H. 2002. Species-specific patterns of litter processing by terrestrial isopods (Isopoda: Oniscidea) in high intertidal salt marshes and coastal forests. Functional Ecology. 16:596-607.

Books and Book Sections

Pennings, S.C. and He, Q. 2021. Community ecology of salt marshes. Pages 82-112 in: Fitzgerald, D. and Hughes, Z. (editors). Salt marshes. Cambridge University Press, New York.

Pennings, S.C. and He, Q. 2021. Community ecology of salt marshes. Pages 82-112 in: Fitzgerald, D. and Hughes, Z. (editors). Salt marshes. Cambridge University Press, New York.

Sharitz, R.R. and Pennings, S.C. 2014. Development of wetland plant communities. Pages 133-150 in: Ecology of Freshwater and Estuarine Wetlands. University of California Press.

Sharitz, R.R., Batzer, D. and Pennings, S.C. 2014. Ecology of Freshwater and Estuarine Wetlands: An Introduction. Pages 1-22 in: Ecology of Freshwater and Estuarine Wetlands. University of California Press.

Pennings, S.C. 2013. Forging collaborations between ecology and historical ecology. Pages 167-175 in: Thompson, V.D. and Waggoner, J. (editors). The archaeology and historical ecology of small scale economies. The University Press of Florida, Gainesville.

Pennings, S.C., Alber, M., Alexander, C.R. Jr., Booth, M.G., Burd, A.B., Cai, W.-J., Craft, C.B., DePratter, C., Di Iorio, D., Hollibaugh, J.T., Hopkinson, C.S., Joye, S.B., Meile, C., Moore, W.S., Silliman, B.R., Thompson, V.D. and Wares, J.P. 2012. South Atlantic Tidal Wetlands. Pages 45-61 in: Batzer, D.P. and Baldwin, A. (editors). Wetland Habitats of North America: Ecology and Conservation Concerns. University of California Press, Berkeley, California.

Sharitz, R.R. and Pennings, S.C. 2006. Development of wetland plant communities. Pages 177-241 in: Batzer, D.P. and Sharitz, R.R. (editors). Ecology of freshwater and estuarine wetlands. University of California Press, Berkeley.

Conference Papers (Peer Reviewed)

Pennings, S.C. 2010. Local and geographic variation in Spartina-herbivore interactions. Pages 9-14 in: Proceedings of the Third International Conference on Invasive Spartina. San Francisco Estuary Invasive Spartina Project of the California State Coastal Conservancy, Oakland, CA.

Conference Posters and Presentations

Adams, T. and Pennings, S.C. Poster: Long-term saltmarsh katydid (Orchelimum fidiciinium) dynamics on Sapelo Island. 103rd Annual meeting of the Ecological Society of America, August 5-10, 2018, New Orleans.

Liu, W. and Pennings, S.C. Presentation: Allometric variation of the salt marsh grass Spartina alterniflora across space and time. 103rd Annual meeting of the Ecological Society of America, August 5-10, 2018, New Orleans.

Pennings, S.C. Presentation: Effects of Hurricanes Matthew and Irma on coastal wetlands in Georgia. 103rd Annual meeting of the Ecological Society of America, August 5-10, 2018, New Orleans.

Zhang, Y. and Pennings, S.C. Presentation: Latitudinal patterns of insect folivory of mangrove forest in southern coast of China. 103rd Annual meeting of the Ecological Society of America, August 5-10, 2018, New Orleans.

Smith, D., Herbert, E., Li, F., Widney, S., Desha, J., Schubauer-Berigan, J.P., Pennings, S.C., Angelini, C., Medeiros, P.M., Byers, J., Alber, M. and Craft, C.B. 2016. Poster: Seawater Addition Long Term Experiment (SALTEx). Georgia Department of Natural Resources Coastal Resources Division 2016 Climate Conference, November 2-3, 2016, Jekyll Island, GA.

Smith, D., Herbert, E., Li, F., Widney, S., Desha, J., Schubauer-Berigan, J.P., Pennings, S.C., Angelini, C., Medeiros, P.M., Byers, J., Alber, M. and Craft, C.B. 2016. Poster: Seawater Addition Long Term Experiment (SALTEx). SEERS 2016 Spring Meeting, March 10-12, 2016, Bluffton, South Carolina.

Gehman, A., McLenaghan, N.A., Byers, J., Alexander, C.R. Jr., Pennings, S.C. and Alber, M. 2015. Poster: Effects of development and shoreline armoring on the high marsh ecosystem. Benthic Society Ecology Meeting 2015, March 4-7, 2015, Quebec City, CN.

Mony, C., Pennings, S.C., Bittebiere, A.-K. and Garbey, M. 2014. Poster: Attack of the clones: multiscale modeling of clonal plant dynamic. 2014 Multi-Scale Modeling Consortium, September 3-4, 2014, NIH campus, Bethesda Maryland.

Pennings, S.C. 2013. Presentation: Computer science meets biology: using volunteer scientists to map salt marsh plant communities. Workshop on complex adaptive systems, 21 October, 2013, La Rochelle, France.

Vu, H. and Pennings, S.C. 2013. Presentation: Biophysical feedbacks in tidal creek formation. Coastal and Estuarine Research Federation 2013, 3-7 November 2013, San Diego, CA.

Garbey, M., Bittebiere, A.-K., Smaoui, M., Rinsurongkawong, W., Clement, B., Pennings, S.C. and Mony, C. 2012. Presentation: Large scale simulations of virtual prairies with volunteer computing. Modeling wetland processes. 2012 LTER All Scientist Meeting, September 10-13, 2012, Estes Park, CO.

Pennings, S.C. 2012. Presentation: Distributed graduate courses in the LTER network. LTER graduate student training: Innovations and cross-site opportunities. 2012 LTER All Scientist Meeting, September 10-13, Estes Park, CO.

Pennings, S.C. 2012. Presentation: Stability of salt marsh plant communities. PDTNet. 2012 LTER All Scientist Meeting, September 10-13, Estes Park, CO.

Wieski, K. and Pennings, S.C. 2012. Poster: Climate drivers of Spartina marsh production on the Georgia coast. LTER All Scientists Meeting, September 10-13, 2012, Estes Park.

Pennings, S.C., Ho, C.-K., Marczak, L., McCall, B.D., Wieski, K. and Vu, H. 2011. Presentation: Geographic variation in salt marsh food webs. Comparative studies of estuarine and coastal system properties. Coastal and Estuarine Research Federation, November 6-11, Daytona Beach, FL.

Vu, H., Wieski, K. and Pennings, S.C. 2011. Presentation: Crab mediated tidal creek formation in salt marshes. Coastal and Estuarine Research Federation, November 6-11, Daytona Beach, FL.

Alexander, C.R. Jr., Alber, M., Hladik, C.M. and Pennings, S.C. 2010. Presentation: Physical-Biological Interactions in Coastal Settings: The Georgia Coastal Ecosystem LTER Example. American Geophysical Union - Meeting of the Americas, 9-13 August 2010, Foz do Iguacu, Brazil.

Guo, H., Pennings, S.C. and Wieski, K. 2010. Presentation: Mechanisms mediating plant distributions in Georgia tidal marshes. 2010 annual meeting of the Ecological Society of America, August 1-6, 2010, Pittsburgh, PA.

Hladik, C.M., Alber, M., Schalles, J.F. and Pennings, S.C. 2010. Poster: Salt marsh habitat mapping on Sapelo Island, GA using LIDAR and hyperspectralimagery. National Estuarine Research Reserve Annual Meeting, October 11-15, 2010, National Conservation Training Center, Shepherdstown, WV.

Ho, C.-K. and Pennings, S.C. 2010. Presentation: Preference and performance in plant-herbivore interactions across latitude. Organized Oral Session, “Latitudinal gradients in consumer-resource interactions: bridging the gap between pattern and process“. 2010 annual meeting of the Ecological Society of America, August 1-6, 2010, Pittsburgh, PA.

Pennings, S.C. 2010. Presentation: Latitudinal variation in plant-herbivore interactions in salt marshes. Organized Oral Session, “Latitudinal gradients in consumer-resource interactions: bridging the gap between pattern and process“. 2010 annual meeting of the Ecological Society of America, August 1-6, 2010, Pittsburgh, PA.

Alber, M., Alexander, C.R. Jr., Hladik, C.M., Jackson, C. and Pennings, S.C. 2009. Presentation: Marsh hammocks: isolated uplands in the marsh environment. Coastal and Estuarine Research Federation meeting, November 2009, Portland, Oregon.

Guo, H., Ewers, C. and Pennings, S.C. 2009. Presentation: Factors mediating the distribution of Suaeda linearis in Georgia coastal marshes. Ecological Society of America 2009 annual meeting, August 2-7, 2009, Albuquerque, NM.

Hladik, C.M., Alber, M., Schalles, J.F., Lynes, A.R. and Pennings, S.C. 2009. Poster: Salt marsh habitat mapping on Sapelo Island, GA using LIDAR and hyperspectral imagery. Coastal and Estuarine Research Federation meeting, November 2009, Portland, Oregon.

Pennings, S.C., Buck, T.L., Lynes, A.R. and Grace, J.B. 2009. Poster: Centrifugal organization of vegetation in salt marsh plant communities. 2009 LTER All Scientists Meeting, September 14-16, 2009, Estes Park, Colorado.

Schalles, J.F., Hladik, C.M., Pennings, S.C., Steele, M.M. and Lynes, A.R. 2009. Presentation: Extracting habitat features from hyperspectral wetland imagery at NOAA’s Sapelo Island and Mission-Aransas National Estuarine Research Reserves. NOAA – Educational Partnership Program Fifth Education and Science Forum, November 13, 2009, Howard University, Washington, D.C.

Guo, H., Pennings, S.C. and Wieski, K. 2008. Poster: Physical stress, plant productivity, competition, and diversity in Georgia tidal marshes. Coastal Habitats. 93rd Annual Meeting of the Ecological Society of America, August 3-8, 2008, Milwaukee, Wisconsin.

Wieski, K., Guo, H. and Pennings, S.C. 2008. Poster: Ecosystem functions of tidal fresh, brackish, and salt marshes. Estuarine, Coastal and Intertidal Systems. 93rd Annual Meeting of the Ecological Society of America, August 3-8, 2008, Milwaukee, Wisconsin.

Collins, S.L., Suding, K.N., Cleland, E.E., Batty, M., Pennings, S.C., Gross, K.L., Grace, J.S., Gough, L., Fargione, J.E., Clark, C.M. Presentation: Rank clocks and plant community dynamics. Joint Ecological Society of America / Society for Ecological Restoration 2007 Meeting, August 5-10, 2007, San Jose, CA.

Ho, C.-K. and Pennings, S.C. 2007. Presentation: Bergmann¡'s Rule and latitudinal variation in herbivore body size. Joint Ecological Society of America / Society for Ecological Restoration 2007 Meeting, August 5-10, 2007, San Jose, California, USA.

Pennings, S.C., Ho, C.-K., Salgado, C., Wieski, K., Dave, N., Kunza, A.E., Wason, E.L. 2007. Presentation: Latitudinal variation in herbivore pressure in Atlantic Coast salt marshes. Joint Ecological Society of America / Society for Ecological Restoration 2007 Meeting, August 5-10, 2007, San Jose, CA.

Volkmer, M., Schalles, J.F., Hladik, C.M. and Pennings, S.C. 2007. Presentation: Coastal wetland and surface water classifications using hyperspectral aerial imagery. CS23 Remote Sensing and Emerging Technologies. American Society of Limnology and Oceanography 2007 Aquatic Science Meetings, February 5, 2007, Sante Fe, N.M.

Wason, E.L. and Pennings, S.C. 2007. Poster: Gradients of grasshopper species composition and size in Atlantic Coast salt marshes. Joint Ecological Society of America / Society for Ecological Restoration 2007 Meeting, August 5-10, 2007, San Jose, CA.

Ho, C.-K. and Pennings, S. C. Presentation: Preference and performance in plant-herbivore interactions across latitude. 91st Annual Meeting of the Ecological Society of America. August 6-11, 2006, Memphis, Tennessee, USA.

Kunza, A.E. and Pennings, S.C. 2006. Poster: Patterns of plant diversity in two salt marsh regions. Long Term Ecological Research Network All-Scientists Meeting, September 20-24, 2006, Estes Park, Colorado.

Pennings, S.C. 2006. Presentation: Plant community response to nitrogen enrichment: results from a cross-site synthesis. Long Term Ecological Research Network All-Scientists Meeting, September 20-24, 2006, Estes Park, Colorado.

Pennings, S.C. 2006. Presentation: Sea-level rise and ecosystem services of tidal marshes. Sea-level rise, hurricanes, and the future of our coasts. Sigma Xi Meeting,Texas A&M University, March 30, 2006.

Clark, C., Cleland, E., Collins, S., Gough, L., Pennings, S. and Suding, K. 2005. Presentation: Predicting the loss of biodiversity with nitrogen addition across North American herbaceous communities: abiotic vs. biotic drivers. Ecological Society of America 2005 Meeting - Ecology at multiple scales, August 7-12, 2005, Montreal, Canada.

Fargione, J., Pennings, S., Clark, C., Suding, K., Gough, L., Grace, J., Cleland, E., Collins, S. and Gross, K. 2005. Presentation: A negative relationship between species richness and productivity response to N addition in a synthesis of N fertilization studies. Ecological Society of America 2005 Meeting - Ecology at multiple scales, August 7-12, 2005, Montreal, Canada.

Ho, C.-K. and Pennings, S.C. 2005. Poster: Consequences of Omnivory for Trophic Interactions: A Case from Salt Marshes. Ecological Society of America 2005 Meeting - Ecology at multiple scales, August 7-12, 2005, Montreal, Canada.

Ho, C.-K. and Pennings, S.C. 2005. Presentation: Consequences of ominivory. The 18th Biennial Conference of the Estuarine Research Federation. Oct. 16-20, 2005, Norfolk, Virginia.

Kunza Vargas, A.E. and Pennings, S.C. 2005. Poster: Plant diversity of Texas and Georgia salt marshes. Ecological Society of America 2005 Meeting - Ecology at multiple scales, August 7-12, 2005, Montreal, Canada.

Pennings, S.C. 2005. Presentation: Physical forcing and variation in salt marsh plant productivity at multiple time scales. Ecological Society of America 2005 Meeting - Ecology at multiple scales, August 7-12, 2005, Montreal, Canada.

Pennings, S.C. 2004. Presentation: Local and geographic variation in Spartina-herbivore interactions. Third International Conference on Invasive Spartina, November 8-10, 2004, San Francisco, CA.

Ewanchuk, P., Pennings, S.C., Bertness, M.D., Selig, E.R. and Houser, L.T. 2003. Presentation: Temporal and spatial variation in interactions among salt marsh plants. Estuaries on the Edge. Estuarine Research Federation and LTER Joint Symposium, Setember 18, 2003, Seattle, WA.

Ho, C.-K., Salgado, C.S., Dave, N.B. and Pennings, S.C. 2003. Poster: Latitudinal variation in herbivore densities and damage to plants in coastal salt marshes. Ecological Society of America, August 3-8, 2003, Savannah, GA.

McFarlin, C.R., Buck, T.L., Newell, S.Y. and Pennings, S.C. 2003. Poster: Impact of fertilization on the abundance of two detritivorous snails. Embarking on a Decade of Synthesis. 2003 LTER All Scientists Meeting, September 18-21, 2003, Seattle, WA.

McFarlin, C.R., Buck, T.L., Newell, S.Y. and Pennings, S.C. 2003. Poster: Impact of fertilization on the abundance of two detritivorous snails. Uplands to Lowlands: Coastal Processes in a Time of Global Change. Aquatic Systems. Ecological Society of America, August 3-8, 2003, Savannah, GA.

Pennings, S.C. 2003. Presentation: Latitudinal variation in plant-herbivore interactions in Atlantic Coast salt marshes. Estuaries on the Edge. Estuarine Research Federation and LTER Joint Symposium, September 18, 2003, Seattle, WA.

Pennings, S.C., Bertness, M.D., Ewanchuk, P.J., Selig, E.R. and Houser, L.T. 2003. Presentation: Temporal and spatial variation in positive and negative interactions among salt marsh plants: the role of climate and community. Expanding scales, expanding perspectives: new insights into marine ecosystem dynamics. Ecological Society of America, August 2003, Savannah, GA.

Salgado, C., Ho, C.-K., Davé, N. and Pennings, S.C. 2003. Poster: Strength of plant-herbivore interactions varies across latitude in Atlantic coast salt marshes. 2003 Meeting of the International Biogeographic Society. International Biogeographic Society, January 4-8, 2003, Mesquite, NV.

Buck, T.L. and Pennings, S.C. 2002. Poster: Variation in zonation patterns of salt marsh plants among sites. 87th Annual Meeting of the Ecological Society of America and the 14th Annual International Conference of the Society for Ecological Restoration. Ecological Society of America, August, 2002, Tucson, Arizona.

Richards, C.L., Pennings, S.C. and Donovan, L.A. 2002. Poster: Community-wide patterns of phenotypic variation in salt marsh plants. 2002 National Estuarine Research Reserve (NERR) Annual Meeting.

Amft, J., Blanton, J.O., Di Iorio, D., Helm, K. and Pennings, S.C. 2001. Poster: Estuarine salinity regime near Sapelo Island, Georgia. ERF 2001: An Estuarine Odyssey. Estuarine Research Federation, November 4-8, 2001, St. Petersburg, Florida.

Goranson, C. and Pennings, S.C. 2001. Poster: Effects of physical gradients on palatability of salt marsh plants to herbivores. Poster presented at the ESA Annual Meeting 2001. Ecological Society of America, Madison, Wisconsin.

Pennings, S.C., Bertness, M.D., Donnelly, J.P., Ewanchuk, P.J., Silliman, B.R. and Callaway, R.M. 2001. Presentation: Impacts of global change on coastal salt marshes. Keynote address to the German Limnological Association, September 17-21, 2001, Kiel, Germany.

Pennings, S.C., Bertness, M.D., Donnelly, J.P., Ewanchuk, P.J., Silliman, B.R. and Callaway, R.M. 2001. Presentation: Impacts of human activities on coastal salt marshes. Meeting of the Western Society of Naturalists, November 10-12, 2001, Ventura, California.

Richards, C.L., Pennings, S.C. and Donovan, L.A. 2000. Presentation: Morphological variation in twelve salt marsh plant species on Sapelo Island, GA. ESA 2000. Ecological Society of America, Aug 01, 2000.