Abstract
The long-term response of riparian communities to shifting climatic conditions can be addressed by the ichnofossil record, because organism behavior is typically altered in response to changes in environmental factors. During the late Paleozoic, the Appalachian Basin experienced a shift from an ever-wet to wet–dry climate. Changes in the abundance, diversity, density, and composition of ichnofossil assemblages were investigated in fluvial point bar sandstones from five roadside outcrops of the Middle Pennsylvanian-to-early Permian Allegheny, Conemaugh, Monongahela, and Dunkard groups located in southeast Ohio and northwest West Virginia. Ichnofossil data were collected using a 0.5 × 0.5-m grid placed on bedding plane surfaces and from vertically oriented thin sections. Abundance, density, diversity, and burrow widths increased through the study interval. Behaviors changed from stationary- to mobile-deposit feeding, while community composition shifted toward more established, permanent generalists. These changes in ichnofossil assemblages suggest that the shift to a drier, more pronounced seasonal climate made short- to long-term occupation of the point bar sands more advantageous as surface conditions were more unfavorable and resources limited. This study helps us understand how terrestrial community composition and ecosystem dynamics shift over long time intervals in response to environmental perturbations. By assessing these changes, we can better predict what future impacts climatic shifts will have on continental ecosystems and terrestrial communities.
Similar content being viewed by others
Availability of data and materials
All data collected are available in the tables and appendices.
Code availability
Statistical analyses were performed using PAST version 3, free software available at https://folk.uio.no/ohammer/past/.
References
Aslan, A., & Autin, W. J. (1998). Holocene flood-plain soil formation in the southern lower Mississippi Valley: Implications for interpreting alluvial paleosols. Geological Society of America Bulletin, 110(4), 433–449.
Beatty, T. W., Zonneveld, J. P., & Henderson, C. M. (2008). Anomalously diverse Early Triassic ichnofossil assemblages in northwest Pangea: A case for a shallow-marine habitable zone. Geology, 36(10), 771–774.
Blakey, R.C. (2008). Gondwana paleogeography from assembly to breakup—a 500 my odyssey. In C. R. Fielding, T. D. Frank, & J. L. Isbell (Eds.), Resolving the Late Paleozoic Ice Age in time and space. Geological Society of America. pp. 1–28
Boyce, C. K., & Lee, J. E. (2010). An exceptional role for flowering plant physiology in the expansion of tropical rainforests and biodiversity. Proceedings. Biological Sciences, 277(1699), 3437–3443. https://doi.org/10.1098/rspb.2010.0485
Brezinski, D. K., & Kollar, A. D. (2011). Pennsylvanian climatic events and their congruent biotic responses in the central Appalachian Basin. Field Guides, 20, 45–60.
Brierley, G. J., Ferguson, R. J., & Woolfe, K. J. (1997). What is a fluvial levee? Sedimentary Geology, 114(1–4), 1–9.
Bromley, R. G. (1996). Trace fossils: Biology, taphonomy and applications. Chapman and Hall.
Buatois, L. A., & Mángano, M. A. (2004). Animal-substrate interactions in freshwater environments: applications of ichnology in facies and sequence stratigraphic analysis of fluvio-lacustrine successions. In D. Mcllroy (Ed.), The application of ichnology to palaeoenvironmental and stratigraphic analysis (Vol. 228, pp. 311–334). Geological Society Special Publication, Geological Society of London.
Buatois, L., & Mángano, M. G. (2011). Ichnology: organism-substrate interactions in space and time. Cambridge University Press.
Buatois, L. A., Mángano, M. G., Genise, J. F., & Taylor, T. N. (1998). The ichnologic record of the continental invertebrate invasion; evolutionary trends in environmental expansion, ecospace utilization, and behavioral complexity. Palaios, 13(3), 217–240.
Cadée, G. C. (1984). ‘Opportunistic feeding’, a serious pitfall in trophic structure analysis of (paleo) faunas. Lethaia, 17(4), 289–292.
Capon, S. J., Chambers, L. E., Mac Nally, R., Naiman, R. J., Davies, P., Marshall, N., Pittock, J., Reid, M., Capon, T., Douglas, M., Catford, J., Baldwin, D. S., Stewardson, M., Roberts, J., Parsons, M., & Williams, S. E. (2013). Riparian ecosystems in the 21st century: Hotspots for climate change adaptation? Ecosystems, 16(3), 359–381.
Catena, A., & Hembree, D. (2012). Recognizing vertical and lateral variability in terrestrial landscapes: A case study from the paleosols of the Late Pennsylvanian Casselman Formation (Conemaugh Group) southeast Ohio, USA. Geosciences, 2(4), 178–202.
Cecil, C. B. (2013). An overview and interpretation of autocyclic and allocyclic processes and the accumulation of strata during the Pennsylvanian–Permian transition in the central Appalachian Basin, USA. International Journal of Coal Geology, 119, 21–31.
Cecil, C. B., Stanton, R. W., Neuzil, S. G., Dulong, F. T., Ruppert, L. F., & Pierce, B. S. (1985). Paleoclimate controls on late Paleozoic sedimentation and peat formation in the central Appalachian Basin (USA). International Journal of Coal Geology, 5(1–2), 195–230.
Cecil, C. B., Brezinski, D. K., & Dulong, F. (2004). The Paleozoic record of changes in global climate and sea level: Central Appalachian Basin (Vol. 1264, pp. 77–133). Geology of the national capital region—Field trip guidebook: US Geological Survey Circular.
Chan, E. K., Yu, Y. T., Zhang, Y., & Dudgeon, D. (2008). Distribution patterns of birds and insect prey in a tropical riparian forest. Biotropica, 40(5), 623–629.
Cloudsley-Thompson, J. L. (1975). Adaptations of Arthropoda to arid environments. Annual Review of Entomology, 20(1), 261–283.
Collinson, J. D. (1996). Alluvial sediments. In H. G. Reading (Ed.), Sedimentary environments: Processes, facies, and stratigraphy (pp. 37–82). Wiley.
Condit, D. D. (1912). The Conemaugh formation in Ohio. Ohio Geological Survey 4th Series. Bulletin, 17, 363.
Condron, L., Stark, C., O’Callaghan, M., Clinton, P., & Huang, Z. (2010). The role of microbial communities in the formation and decomposition of soil organic matter. In G. R. Dixon & E. L. Tilston (Eds.), Soil microbiology and sustainable crop production (pp. 81–118). Springer.
Crowell, J. C. (1995). The ending of the late Paleozoic ice age during the Permian Period. In P. A. Scholle, T. M. Peryt, & D. S. Ulmer-Scholle (Eds.), The Permian of Northern Pangea (pp. 62–74). Springer.
DiMichele, W. A. (2014). Wetland-dryland vegetational dynamics in the Pennsylvanian ice age tropics. International Journal of Plant Sciences, 175(2), 123–164.
DiMichele, W. A., Pfefferkorn, H. W., & Phillips, T. L. (1996). Persistence of Late Carboniferous tropical vegetation during glacially driven climatic and sea-level fluctuations. Palaeogeography, Palaeoclimatology, Palaeoecology, 125, 105–128.
DiMichele, W. A., Montañez, I. P., Poulsen, C. J., & Tabor, N. J. (2009). Climate and vegetational regime shifts in the late Paleozoic ice age earth. Geobiology, 7(2), 200–226.
Dimmitt, M. A. (2000). Biomes and communities of the Sonoran Desert Region. In S. J. Phillips & P. WentworthComus (Eds.), A natural history of the Sonoran Desert: Tucson, AZ, US, Arizona-Sonora Desert (pp. 3–18). Museum Press.
Dunkard Group, Pennsylvania-West Virginia-Ohio, USA. Journal of Coal Geology, 119, 79–87.
Dzenowski, N. D., & Hembree, D. I. (2014). The neoichnology of two terrestrial ambystomatid salamanders: Quantifying amphibian burrows using modern analogs. In D. I. Hembree, B. F. Platt, & J. J. Smith (Eds.), Experimental approaches to understanding fossil organisms (pp. 305–341). Springer.
Ekdale, A. A. (1985). Paleoecology of the marine endobenthos. Palaeogeography, Palaeoclimatology, Palaeoecology, 50(1), 63–81.
Erwin, D. H. (2009). Climate as a driver of evolutionary change. Current Biology, 19(14), R575–R583.
Fedorko, N., & Skema, V. (2013). A review of the stratigraphy and stratigraphic nomenclature of the Dunkard Group in West Virginia and Pennsylvania, USA. International Journal of Coal Geology, 119, 2–20.
Ferm, J. C. (1970). Allegheny deltaic deposits. In J. P. Morgan & R. H. Shaver (Eds.), Deltaic sedimentation, modern and ancient (Vol. 15, pp. 246–255). SEPM Special Publication Society of Economic Paleontologists and Mineralogists. https://doi.org/10.2110/pec.70.11
Frey, R. W., Pemberton, S. G., & Fagerstrom, J. A. (1984). Morphological, ethological, and environmental significance of the ichnogenera Scoyenia and Ancorichnus. Journal of Paleontology, 58(2), 511–528.
Garcés, B. L. V., Gierlowski-Kordesch, E., & Bragonier, W. A. (1997). Pennsylvanian continental cyclothem development: No evidence of direct climatic control in the Upper Freeport Formation (Allegheny Group) of Pennsylvania (northern Appalachian Basin). Sedimentary Geology, 109(3–4), 305–319.
Gastaldo, R. A., DiMichele, W. A., & Pfefferkorn, H. W. (1996). Out of the icehouse into the greenhouse: a late Paleozoic analog for modern global vegetational change. GSA Today, 6, 1–7.
Getty, P. R., McCarthy, T. D., Hsieh, S., & Bush, A. M. (2016). A new reconstruction of continental Treptichnus based on exceptionally preserved material from the Jurassic of Massachusetts. Journal of Paleontology, 90(2), 269–278.
Gingras, M. K., Bann, K. L., MacEachern, J. A., Waldron, J., & Pemberton, S. G. (2007). A conceptual framework for the application of trace fossils. In J. A. MacEachern, K. L. Bann, M. K. Gingras, & S. G. Pemberton (Eds.), Applied ichnology. SEPM short course notes (Vol. 52, pp. 1–26). Society for Sedimentary Geology.
Häntzschel, W. (1975). Treatise on invertebrate paleontology. In P. W. Miscellanea (Ed.), Supplement 1. Trace fossils and problematica (2nd ed.). The Geological Society of America.
Hasiotis, S. T. (2002). Continental trace fossils. SEPM Short Course Notes (Vol. 51). Society for Sedimentary Geology.
Hasiotis, S. T. (2004). Reconnaissance of Upper Jurassic Morrison Formation ichnofossils, Rocky Mountain Region, USA: Paleoenvironmental, stratigraphic, and paleoclimatic significance of terrestrial and freshwater ichnocoenoses. Sedimentary Geology, 167(3–4), 177–268.
Hasiotis, S. T., & Dubiel, R. F. (1993). Continental trace fossils of the Upper Triassic Chinle Formation, Petrified Forest National Park, Arizona. The nonmarine Triassic. Bulletin of the New Mexico Museum of Natural History and Science, 3, 175–178.
Hasiotis, S. T., Dubiel, R. F., & Demko, T. M. (1998). A holistic approach to reconstructing Triassic paleoecosystems: Using ichnofossils and paleosols as a basic framework. National Park Service Paleontological Research, 3, 122–124.
Hasiotis, S. T., Kraus, M. J., & Demko, T. M. (2007). Climatic controls on continental trace fossils. In W. Miller (Ed.), Trace fossils: Concepts, problems, prospects (pp. 172–195). Elsevier.
Hembree, D. I. (2009). Neoichnology of burrowing millipedes: linking modern burrow morphology, organism behavior, and sediment properties to interpret continental ichnofossils. Palaios, 24, 425–439. https://doi.org/10.2110/palo.2008.p08-098r
Hembree, D. I. (2017). Neoichnology of tarantulas (Araneae: Theraphosidae): criteria for recognizing spider burrows in the fossil record. Palaeontologia Electronica. https://doi.org/10.26879/780
Hembree, D. (2018). The role of continental trace fossils in Cenozoic paleoenvironmental and paleoecological reconstructions. In D. A. Croft, D. F. Su, & S. W. Simpson (Eds.), Methods in paleoecology: Reconstructing Cenozoic terrestrial environments and ecological communities (pp. 185–214). Springer. https://doi.org/10.1007/978-3-319-94265-0
Hembree, D. I. (2019). Burrows and ichnofabric produced by centipedes: modern and ancient examples. Palaios, 34, 468–489. https://doi.org/10.2110/palo.2019.059
Hembree, D. I. (2022). Early effects of the Late Paleozoic climate transition on soil ecosystems of the Appalachian Basin (Conemaugh, Monongahela, and Dunkard groups): Evidence from ichnofossils. Palaios, 37(11), 671–690. https://doi.org/10.2110/palo.2021.071
Hembree, D. I., & Blair, M. G. (2016). A paleopedological and ichnological approach to interpreting spatial and temporal variability in Early Permian fluvial deposits of the lower Dunkard Group, West Virginia, USA. Palaeogeography, Palaeoclimatology, Palaeoecology, 454, 246–266.
Hembree, D. I., & Bowen, J. J. (2017). Paleosols and ichnofossils of the Upper Pennsylvanian-Lower Permian Monongahela and Dunkard groups (Ohio, USA): A multi-proxy approach to unraveling complex variability in ancient terrestrial landscapes. Palaios, 32, 295–320.
Hembree, D. I., & Carnes, J. L. (2018). Response of soils and soil ecosystems to the Pennsylvanian–Permian climate transition in the upper fluvial plain of the Dunkard Basin, southeastern Ohio, USA. Geosciences, 8, 203.
Hembree, D. I., & McFadden, C. J. (2020). Analysis of climate and landscape change through the Pennsylvanian and Permian Monongahela and Dunkard Groups, Southeastern Ohio, USA. Journal of Sedimentary Environments, 5, 321–353.
Hembree, D. I., & Nadon, G. C. (2011). A paleopedologic and ichnologic perspective of the terrestrial Pennsylvanian landscape in the distal Appalachian Basin, USA. Palaeogeography, Palaeoclimatology, Palaeoecology, 312, 138–166.
Hembree, D. I., Smith, J. J., Buynevich, I. V., & Platt, B. F. (2017). Neoichnology of semiarid environments: Soils and burrowing animals of the Sonoran Desert, Arizona, USA. Palaios, 32(9), 620–638.
Hils, J. M., & Hembree, D. I. (2015). Neoichnology of the burrowing spiders Gorgyrella inermis (Mygalomorphae: Idiopidae) and Hogna lenta (Araneomorphae: Lycosidae). Palaeontologica Electronica. https://doi.org/10.26879/500
Joeckel, R. M. (1995). Paleosols below the Ames Marine Unit (Upper Pennsylvanian, Conemaugh Group) in the Appalachian Basin, USA: Variability on an ancient depositional landscape. Journal of Sedimentary Research, 65(2a), 393–407.
Jumars, P. A., & Wheatcroft, R. A. (1989). Responses of benthos to changing food quality and quantity, with a focus on deposit feeding and bioturbation. In W. H. Berger, V. S. Smetacek, & G. Wefer (Eds.), Productivity of the ocean: Present and past (pp. 235–253). Wiley.
Kale, V. S., Patil, S. S., Satoskar, V., & Kumar, P. (1997). Occurrence of Planolites from the Nagarjuna Sagar Area, Northwestern Cuddapah Basin. Journal of Geological Society of India, 49(5), 589–596.
Kondolf, G. M., Kattelmann, R., Embury, M., & Erman, D. C. (1996). Status of riparian habitat. In C. I. Millar (Ed.), Sierra Nevada Ecosystem Project: Final report to Congress (pp. 1009–1030). University of California.
Koy, K., & Plotnick, R. E. (2007). Theoretical and experimental ichnology of mobile foraging. In W. Miller (Ed.), Trace fossils: Concepts, problems, prospects (pp. 428–441). Elsevier.
Lavelle, P., & Spain, A. V. (2005). Soil ecology. Springer Publishing.
Lawrence, J. F., & Newton, A. F., Jr. (1982). Evolution and classification of beetles. Annual Review of Ecology and Systematics, 13(1), 261–290.
LeRoux, P. C., & McGeoch, M. A. (2008). Rapid range expansion and community reorganization in response to warming. Global Change Biology, 14(12), 2950–2962.
Lopez, G. R. & Levinton, J. S. (1987). Ecology of deposit-feeding animals in marine sediments. The Quarterly Review of Biology, 62(3), 235–260. http://www.jstor.org/stable/2828974
Lucas, S. G. (2013). Vertebrate biostratigraphy and biochronology of the upper Paleozoic Dunkard Group, Pennsylvania-West Virginia-Ohio, USA. Journal of Coal Geology, 119, 79–87.
MacEachern, J. A., Pemberton, S. G., Bann, K. L., & Gingras, M. K. (2007). Departures from the archetypal ichnofacies: effective recognition of physico-chemical stresses in the rock record. In J. A. MacEachern, K. L. Bann, M. K. Gingras, & S. G. Pemberton (Eds.), Applied ichnology SEPM Short Course Notes (Vol. 52, pp. 65–93). Society for Sedimentary Geology.
Martin, W. D. (1998). Geology of the Dunkard Group (Upper Pennsylvanian-Lower Permian) in Ohio, West Virginia, and Pennsylvania. Ohio Division of Geological Survey Bulletin, 73, 1–49.
Miall, A. D. (2010). Alluvial deposits. In N. P. James & R. W. Dalrymple (Eds.), Facies models 4 (pp. 105–137). Geological Association of Canada.
Mikuś, P., & Uchman, A. (2013). Beetle burrows with a terminal chamber: a contribution to the knowledge of the trace fossil Macanopsis in continental sediments. Palaios, 28, 403–413. https://doi.org/10.2110/palo.2012.p12-129r
Milici, R. C. (2005). Appalachian coal assessment: defining the coal systems of the Appalachian basin. In P. D. Warwick (Ed.), Coal systems (Vol. 387, pp. 9–30). Geological Society of America Special Paper. Geological Society of America.
Miller, M. F. (2003). Styles of behavioral complexity recorded by selected trace fossils. Palaeogeography, Palaeoclimatology, Palaeoecology, 192(1–4), 33–43.
Miller, M. F., & Collinson, J. W. (1994). Trace fossils from Permian and Triassic sandy braided stream deposits, central Transantarctic Mountains. Palaios, 9(6), 605–610.
Minter, N. J., Buatois, L. A., Mángano, M. G., Davies, N. S., Gibling, M. R., & Labandeira, C. (2016). The establishment of continental ecosystems. In M. G. Mángano & L. A. Buatois (Eds.), The trace-fossil record of major evolutionary events: Mesozoic and Cenozoic (Vol. 2, pp. 205–324). Springer.
Montañez, I. P., & Cecil, C. B. (2013). Paleoenvironmental clues archived in non-marine Pennsylvanian–lower Permian limestones of the Central Appalachian Basin, USA. International Journal of Coal Geology, 119, 41–55.
Montañez, I. P., & Poulsen, C. J. (2013). The Late Paleozoic ice age: An evolving paradigm. Annual Review of Earth and Planetary Sciences, 41, 629–656.
Moore, J. C., Berlow, E. L., Coleman, D. C., de Ruiter, P. C., Dong, Q., Hastings, A., Collins Johnson, N., McCann, K. S., Melville, K., Morin, P. J., Nadelhoffer, K., Rosemond, A. D., Post, D. M., Sabo, J. L., Scow, K. M., Vanni, M. J., & Wall, D. H. (2004). Detritus, trophic dynamics and biodiversity. Ecology Letters, 7(7), 584–600.
Morrissey, L. B., & Braddy, S. J. (2004). Terrestrial trace fossils from the Lower Old Red sandstone, southwest Wales. Geological Journal, 39(3–4), 315–336.
Naiman, R. J., Decamps, H., & Pollock, M. (1993). The Role of riparian corridors in maintaining regional biodiversity. Ecological Applications, 3(2), 209–212. https://doi.org/10.2307/1941822
Nakamura, A., Proctor, H., & Catterall, C. P. (2003). Using soil and litter arthropods to assess the state of rainforest restoration. Ecological Management & Restoration, 4, S20–S28.
Pianka, E. R. (1970). On r-and K-selection. The American Naturalist, 104(940), 592–597.
Powell, M. G., Schöne, B. R., & Jacob, D. E. (2009). Tropical marine climate during the late Paleozoic ice age using trace element analyses of brachiopods. Palaeogeography, Palaeoclimatology, Palaeoecology, 280(1–2), 143–149.
Retallack, G. J. (2001). Soils of the past: An introduction to paleopedology. John Wiley and Sons.
Rindsberg, A. K., & Kopaska-Merkel, D. C. (2005). Treptichnus and Arenicolites from the Steven C. Minkin Paleozoic footprint site (Langsettian, Alabama, USA). In R. J. Buta (Ed.), Pennsylvanian footprints in the Black Warrior Basin of Alabama (pp. 121–141). University of Alabama.
Roy, K., Valentine, J. W., Jablonski, D., & Kidwell, S. M. (1996). Scales of climatic variability and time averaging in Pleistocene biotas: Implications for ecology and evolution. Trends in Ecology & Evolution, 11(11), 458–463.
Schneider, R. J., Lucas, S. G., & Barrick, J. E. (2013). The Early Permian age of the Dunkard Group, Appalachian basin, U.S.A., based on spiloblattinid insect biostratigraphy. International Journal of Coal Geology, 119, 88–92.
Stow, D. A. V. (2005). Sedimentary rocks in the field: a color guide. Gulf Professional Publishing.
Sturgeon, M. T. (1958). The geology and mineral resources of Athens County, Ohio. Ohio Division of Geological Survey Bulletin, 57, 1–600.
Sturgeon, M. T., & Merrill, W. M. (1949). An additional fossiliferous member in the Allegheny formation (Pennsylvanian) of Ohio. The Ohio Journal of Science, 49(1), 1–11.
Tabor, N. J., & Montañez, I. P. (2004). Morphology and distribution of fossil soils in the Permo-Pennsylvanian Wichita and Bowie Groups, north-central Texas, USA: Implications for western equatorial Pangean palaeoclimate during icehouse–greenhouse transition. Sedimentology, 51(4), 851–884.
Tabor, N. J., & Poulsen, C. J. (2008). Palaeoclimate across the Late Pennsylvanian-Early Permian tropical palaeolatitudes: A review of climate indicators, their distribution, and relation to palaeophysiographic climate factors. Palaeogeography, Palaeoclimatology, Palaeoecology, 268, 293–310.
Tabor, N. J., Montañez, I. P., Scotese, C. R., Poulsen, C. J., & Mack, G. H. (2008). Paleosol archives of environmental and climatic history in paleotropical western Pangea during the latest Pennsylvanian through Early Permian. In C. R. Fielding, T. D. Frank, & J. L. Isbell (Eds.), Resolving the Late Paleozoic Ice Age in time and space (pp. 291–304). Geological Society of America.
Thacker, H. A., & Hembree, D. I. (2021). Neoichnological study of burrowing darkling beetles (Coleoptera: Tenebrionidae) from larval to adult stages. Ichnos, 28(4), 290–308.
Underwood, E. C., Olson, D., Hollander, A. D., & Quinn, J. F. (2014). Ever-wet tropical forests as biodiversity refuges. Nature Climate Change, 4(9), 740–741.
Vossler, S. M., & Pemberton, S. G. (1988). Skolithos in the Upper Cretaceous Cardium Formation: An ichnofossil example of opportunistic ecology. Lethaia, 21(4), 351–362.
Walsh, R. P., Blake, W. H., Slaymaker, O., & Spencer, T. (2009). Tropical rainforests. In O. Slaymaker, T. Spencer, & C. Embleton-Hamnn (Eds.), Geomorphology and global environmental change (pp. 214–257). Cambridge University Press.
Walther, G. R. (2010). Community and ecosystem responses to recent climate change. Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences, 365, 2019–2024. https://doi.org/10.1098/rstb.2010.0021
Whittaker, R. H. (1953). A consideration of climax theory: The climax as a population and pattern. Ecological Monographs, 23(1), 41–78.
Williams, E. G. (1960). Marine and fresh water fossiliferous beds in the Pottsville and Allegheny Groups of western Pennsylvania. Journal of Paleontology, 34, 908–922.
Zhao, Z., Fan, R. Y., Zhang, L. J., Rodríguez-Tovar, F. J., & Gong, Y. M. (2020). Behavioural responses of Rhizocorallium to storm events: Evidence from the Middle Triassic of SW China. Palaeogeography, Palaeoclimatology, Palaeoecology, 545, 109640.
Zonneveld, J. P., Pemberton, S. G., Saunders, T. D., & Pickerill, R. K. (2002). Large, robust Cruziana from the Middle Triassic of northeastern British Columbia: Ethologic, biostratigraphic, and paleobiologic significance. Palaios, 17(5), 435–448.
Acknowledgements
We would like to thank Joseph Wislocki for help with field work. This project would not have been possible without funding by the Ohio University Geological Sciences Graduate Student Alumni Grant (to JKC), the Geological Society of America Student Research Grant (to JKC), the Society for Sedimentary Geology Student Research Grant (to JKC), and the Paleontological Society Student Research Grant (to JKC).
Funding
Funding for this research was provided by the Ohio University Geological Sciences Graduate Student Alumni Grant (to JKC), the Geological Society of America Student Research Grant (to JKC), the Society for Sedimentary Geology Student Research Grant (to JKC), and the Paleontological Society Student Research Grant (to JKC).
Author information
Authors and Affiliations
Contributions
J.C. and D.H. wrote the manuscript text and prepared the figures and tables. All authors reviewed the manuscript.
Corresponding author
Ethics declarations
Conflict of interest
The authors have no competing interests to declare that are relevant to the content of this article.
Additional information
Communicated by M. V. Alves Martins
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Crowell, J.K., Hembree, D.I. Climate-induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian Appalachian Basin. J. Sediment. Environ. 8, 261–282 (2023). https://doi.org/10.1007/s43217-023-00132-y
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s43217-023-00132-y