Patterns of Variation in the Seed Morphology of Iva annua var. macrocarpa, an Extinct North American Domesticate

  • Andrew W. Weiland The Ohio State University, Columbus, OH.
  • Kristen J. Gremillion The Ohio State University, Columbus, OH.
Keywords: Paleoethnobotany, Seed morphology, Archaeology, Native North America, Domestication


Using measurements from archaeological achenes of the extinct North American cultigen marshelder (Iva annua var. macrocarpa [S.F. Blake] R.C. Jackson), we quantitatively explore patterns of variation of fruit length and width across mid-continental North America. Linear regression shows that while achene length and width increase significantly over time (length: p-value<0.0001, b=-126.04, r2=0.1037, width: p-value<0.0001, b=-230.85, r2=0.0964), overall, regions tend to show more variation. A high incidence of phenotypic variation among domesticated marshelder as measured by coefficient of variation may be a result of introgression with wild stands. An ANOVA Tukey post-hoc analysis of archaeological site samples resulted in homogeneous subsets which correspond to region with some overlap, interpreted as a cline. These results and the low numbers of wild-sized achenes in archaeological marshelder samples of eastern Kentucky support human introduction of domesticated marshelder into this region. Marshelder in the archaeological record reflects the long-standing mixed economies of hunting-gathering and agriculture used by indigenous communities of eastern North America.

Author Biographies

Andrew W. Weiland, The Ohio State University, Columbus, OH.
Andrew W. Weiland is a PhD Candidate in the Department of Anthropology at The Ohio State University in Columbus, OH.
Kristen J. Gremillion, The Ohio State University, Columbus, OH.
Kristen J. Gremillion is a professor in the Department of Anthropology at The Ohio State University in Columbus, OH.


Allaby, R. 2010. Integrating the Processes in the Evolutionary System of Domestication. Journal of Experimental Botany 61:935–944. DOI:10.1093/jxb/erp382.

Anderson, E. 1956. Man as a Maker of New Plants and New Plant Communities. In Man's Role in Changing the Face of the Earth, edited by W. L. Thomas, pp. 763–777. University of Chicago Press, Chicago, IL.

Asch, D.L. and N.B. Asch. 1978. The Economic Potential of Iva annua and its Prehistoric Importance in the Lower Illinois Valley. In The Nature and Status of Ethnobotany, edited by R. I. Ford, pp. 300–346. Museum of Anthropology Anthropological Papers, University of Michigan, Ann Arbor, MI.

Asch, D.L. and N.B. Asch. 1985. Prehistoric Plant Cultivation in West-Central Illinois. In Prehistoric Food Production in North America, edited by R. I. Ford, pp. 149–204. Museum of Anthropology Anthropological Papers, University of Michigan, Ann Arbor, MI.

Black, M. 1963. The Distribution and Archaeological Significance of the Marshelder Iva annua L. Papers of the Michigan Academy of Science, Arts and Letters XLVIII:541–547.

Blake, S.F. 1939. A New Variety of Iva ciliata from Indian Rock Shelters in the South-Central United States. Rhodora 41:81–86.

Chapman, J. and A.B. Shea. 1981. The Archaeobotanical Record: Early Archaic Period to Contact in the Lower Little Tennessee River Valley. Tennessee Anthropologist 6:62–84.

Cowan, C.W. 1979. Excavations at the Haystack Rockshelters, Powell County, Kentucky. Midcontinental Journal of Archaeology 4:3–33. DOI:10.2307/20707828.

Cowan, C.W. 1985. Understanding the Evolution of Plant Husbandry in Eastern North America: Lessons from Botany, Ethnography, and Archaeology. In Prehistoric Food Production in North America, edited by R. I. Ford, pp. 205–243. Museum of Anthropology Anthropological Papers, University of Michigan, University of Michigan, Ann Arbor, MI.

Crites, G.D. 1993. Domesticated Sunflower in Fifth Millenium BP Temporal Context: New Evidence from Middle Tennessee. American Antiquity 58:146–148. DOI:10.2307/281459.

Faulkner, C.T., and G.M. Crothers. 1991. Morphological Variation in Iva annua Achenes: A Statistical Analysis of Archaeological Collections from Eastern North America. Paper presented at the 14th Annual Meeting of the Society of Ethnobiology, St. Louis, MO.

Fritz, G.J. 1986. Prehistoric Ozark Agriculture: The University of Arkansas Rockshelter Collections. Doctoral Dissertation, Department of Anthropology, University of North Carolina at Chapel Hill, Chapel Hill, NC.

Fuller, D.Q. 2007. Contrasting Patterns in Crop Domestication and Domestication Rates: Recent Archaeological Insights from the Old World. Annals of Botany 100:903–924. DOI:10.1093/aob/mcm048.

Fuller, D.Q., E. Asouti and M.D. Purugganan. 2012. Cultivation as Slow Evolutionary Entanglement: Comparative Data on Rate and Sequence of Domestication. Vegetation History and Archaeobotany 21:131–145. DOI:10.1007/s00334-011-0329-8.

Gilmore, M. 1931. Vegetal Remains of the Ozark Bluff-Dweller Culture. In Michigan Academy of Science Arts and Letters14. University of Michigan, Ann Arbor, MI.

Gremillion, K.J. 1993. The Evolution of Seed Morphology in Domesticated Chenopodium: An Archaeological Case Study. Journal of Ethnobiology 13:149–169.

Gremillion, K.J. 1996. Early Agricultural Diet in Eastern North America: Evidence from Two Kentucky Rockshelters. American Antiquity 61:520-536. DOI:10.2307/281837.

Gremillion, K.J. 1997. New Perspectives on the Paleoethnobotany of the Newt Kash Shelter. In People, Plants, and Landscapes, edited by K. J. Gremillion, pp. 23–41. University of Alabama Press, Tuscaloosa, AL.

Gremillion, K.J. 2004. Seed Processing and the Origins of Food Production in Eastern North America. American Antiquity 69:215–233. DOI:10.2307/4128417.

Harlan, J.R., J.M.J. de Wet, and E.G. Price. 1973. Comparative Evolution of Cereals. Evolution 27:311–325. DOI:10.2307/2406971.

Harris, D.R. 1990. Vavilov's Concept of Centres of Origin of Cultivated Plants: Its Genesis and Its Influence on the Study of Agricultural Origins. Biological Journal of the Linnean Society 39:7–16. DOI:10.1111/j.1095-8312.1990.tb01608.x.A.

Jackson, R.C. 1960. A Revision of the Genus Iva L. The University of Kansas Science Bulletin XLI:793–875.

Jones, V.H. 1936. The Vegetal Remains of Newt Kash Hollow Shelter. In Rockshelters in Menifee County, Kentucky, edited by W.S. Webb and W.D. Funkhauser, pp. 147–165. University of Kentucky Reports in Anthropology and Archaeology, Lexington, KY.

Leishmann, M R., I.J. Wright, A.T. Moles, and M. Westboy. 2001. The Evolutionary Ecology of Seed Size. In Seeds: The Ecology of Regeneration in Plant Communities, 2nd edition, edited by M. Fenner, pp. 31-50. Oxford University Press, Cabi Pub, Wallingford, UK.

Linton, R. 1924. The Significance of Certain Traits in North American Maize Culture. American Anthropologist 26:345–349. DOI: 10.1525/aa.1924.26.3.02a00050.

Mueller, N.G. 2017 Documenting Domestication in a Lost Crop (Polygonum erectum L.): Evolutionary Bet-hedgers Under Cultivation. Vegetation History and Archaeobotany 1:1–15. DOI:10.1007/S00334-016-0592-9.

Mueller, N.G., G.J. Fritz, P. Patton, S. Carmody, and E.T. Horton. 2017. Growing the Lost Crops of Eastern North America's Original Agricultural System. Nature Plants 3:7. DOI:10.1038/nplants.2017.92.

Nassaney, M.S., N.H. Lopinot, B.M. Butler, and R.W. Jefferies. 1983. The 1982 Excavations at the Cahokia Interpretive Center Tract, St. Clair County, Illinois. Illinois Department of Conservation Division of Historic Sites.

National Geographic, i-cubed. 2013. Topographic Map [basemap]. Scale Not Given. USGS Topographic Maps. Available at: Accessed May 16, 2018.

Ramsey, C.B., M. Dee, S. Lee, T. Nakagaw, and R. Staff. 2010. Developments in the Calibration and Modelling of Radiocarbon Dates. Radiocarbon 52:953–961. DOI:10.1017/S0033822200046063.

Smith, B.D. 1985. The Role of Chenopodium as a Domesticate in Pre-Maize Garden Systems of the Eastern United States. Southeastern Archaeology 4:51–72.

Smith, B.D. 2001. Low-Level Food Production. Journal of Archaeological Research 9:1-43. DOI:10.1023/A:1009436110049.

Smith, B.D. 2006 Documenting Domesticated Plants in the Archaeological Record. In Documenting Domestication: New Genetic and Archaeological Paradigms, edited by M.A. Zeder, D.G. Bradley, E. Emshwiller, and B.D. Smith, pp. 15–24. University of California Press, Berkeley, CA.

Smith, B.D. 2011. The Cultural Context of Plant Domestication in Eastern North America. Current Anthropology Supplement:S471-S484. DOI:10.1086/659645.

Smith, B.D., C.W. Cowan and M.P. Hoffman. 1992. Rivers of Change: Essays on Early Agriculture in Eastern North America. Smithsonian Institution Press, Washington, DC.

Smith, B.D. and R.A. Yarnell. 2009. Initial Formation of an Indigenous Crop Complex in Eastern North America at 3800 BP. Proceedings of the National Academy of Sciences 106:6561–6566. DOI:10.1073/pnas.0901846106

Stafford, B.D., M.B. Sant, and D.L. Asch. 1985. Smiling Dan: Structure and Function at a Middle Woodland Settlement in the Lower Illinois Valley. Published for the Illinois Dept. of Transportation by the Center for American Archeology, Kampsville Archeological Center, Kampsville, IL.

Steel, R.G.D. and J.H. Torrie. 1980. Principles and Procedures of Statistics: A Biometrical Approach. McGraw-Hill, New York, NY.

USDA, NRCS. 2017. The PLANTS Database. National Plant Data Team, Greensboro, NC 27401–4901 USA. Available at: Accessed December 12, 2017.

Verill, S. 2013. Exact Confidence Bounds for a Normal Distribution Coefficient of Variation. Forestry Service, Department of Agriculture. Available at Accessed December 12, 2017.

Verrill, S. and R.A. Johnson. 2007. Confidence Bounds and Hypothesis Tests for Normal Distribution Coefficients of Variation. Communications in Statistics: Theory and Methods 36:2187–2206. DOI:10.1080/03610920701215126.

Weiland, A.W. 2013. Marshelder (Iva annua L.) Seed Morphology and Patterns of Domestication in Eastern North America. Master’s Thesis, Department of Anthropology, The Ohio State University, Columbus, OH.

Woodburn, J. 1982. Egalitarian Societies. Man 17:431–451. DOI:10.2307/2801707.

Yarnell, R.A. 1972. Iva annua var. macrocarpa: Extinct American Cultigen? American Anthropologist 74:335–341.

Yarnell, R.A. 1981. Inferred Dating of Ozark Bluff Dweller Occupations Based on Achene Size of Sunflower and Sumpweed. Journal of Ethnobiology 1:55–60.

How to Cite
Weiland, A. W., & Gremillion, K. J. (2018). Patterns of Variation in the Seed Morphology of Iva annua var. macrocarpa, an Extinct North American Domesticate. Ethnobiology Letters, 9(2), 75-89.
Research Communications