iTHE HISTORY,  TAXONOMIC STATUS, AND NUTRITIONAL 
COMPONENTS OF THE PREHISTORIC AMERICAN INDIAN FOOD SEED PLANT 
IVA ANNUA 
L. ByPeter Howie CarringtonA DISSERTATION
Submitted toMichigan State Universityfor the degree ofPlant Biology ΠDoctor of Philosophy2015iiABSTRACTTHE HISTORY,  TAXONOMIC STATUS, AND NUTRITIONAL 
COMPONENTS OF THE PREHISTORIC AMERICAN INDIAN FOOD SEED PLANT 
IVA ANNUA 
L. ByPeter Howie CarringtonThe taxonomic status of the plant species Iva annua, an ancient food seed domesticated by American 
-species currently recognized are in fact valid. Metrics of the morphology demonstrate that the two extant subspecies should be treated as synonymous, leaving only 1 valid, existing subspecies, and one from the archaeological history of its domestication. The nutritional properties of the oil and its 
fatty acids, and protein and its amino acids were investigated. Qualitative analysis of the fatty acids showed the composition to be comparable to two other oilseed crops in the Asteraceae. The major
-ity fatty acid is the 18:2 n-6 which is present at approximately three times the levels of the 18:1 fatty acid. Protein analysis, while showing that I. annua Helianthus ann-uus and Carthamus tinctorius, also showed kernel protein levels considerably higher than previously reported for this species, roughly in the range of highs of more than 60 percent. These high nutrient 
levels mean that the contribution of Iva annua to the diet of the ancient Native Americans makes the 
enigma of its abandonment even more profound.iiiCopyright byPETER HOWIE CARRINGTON
2015ivsupportive wife Linda, and to my Uncle Charles Kemp Carrington, both of whom have shaped my life in ways brilliant and unfathomable.vACKNOWLEDGMENTS I would like to thank the late Charles Heiser Jr. for his inspiration and advice and his gift of 
Iva an-nua seed. I would like to thank my committee members; chair Frank Telewski, Rebecca Grumet, Yair 
Shachar-Hill, and Catherine Yansa for the extensive help and patience they have offered and showed 
during the course of this project. I also thank Lisa Carey, Michael Jurgens, Michael Opperman, and 
Mike Pollard for their assistance in obtaining GC-FID and NMR results during lipid analysis; Jameel Al-Haddad for help in statistics and many things. Thanks also to Robert Kaul at the University of 

New Lives for Ancient 
and Extinct Crop
viTABLE OF CONTENTS
LIST OF TABLES
 viiLIST OF FIGURES
 viiiCHAPTER 1 ΠThe Biology and History of Marshelder, 
Iva annua 1  Introduction
 1  Description 1  Archaeology
 4  
Iva Cypselae Over Time
 11
  Measuring Iva annua Cypselae 13 Minerals and Vitamins
 18 Research Question 1
 21 Research Question 2
 21 Research Question 3
 21 Research Question 4
 21 REFERENCES 24CHAPTER 2 ΠA Short Taxonomic History of Genus 
Iva (ASTERACEAE) 21 REFERENCES 45CHAPTER 3 Π The Taxonomic History of 
Iva annua, (ASTERACEAE) 49  
Abstract 49   
Introduction
 49  
Pre-Linnaean & Linnaean History
 50
  
Archaeological Discovery
 65   Iva annua var. 
annua Diagnosis 67  
Conclusion 70  REFERENCES 71CHAPTER 4 ΠA revision of the taxonomy of 
Iva annua to place the variety  75 
                           Iva annua caudata in synonymy with I. a. annua  
Materials and Methods 76  
Results and Discussion 77  
Conclusion 78  
REFERENCES 80CHAPTER 5 ΠProtein and Lipid Content and Composition of 
 82  
              Modern Populations of Iva annua  
Introduction
 82   
Protein 83   
Lipids 86   Materials and Methods 87   
Plant Material 87 vii
   
Protein Analysis
 88   
Lipid Analysis
 89    
Germination Tests
 91  
Results 92   
Total Protein
 92   
Amino Acids 92   
Total Lipids
 94    
Germination Tests
 98  
Discussion 100   
Total Protein
 100   
Amino Acids 101   
Lipids 104  
Conclusions 111
  REFERENCES 116
  viii
LIST OF TABLES
Table 1-1
 Selected Mineral and Vitamin Content (mg/100g edible portion) of
 Three Asteraceae Seeds Grown as Crops
 19Table 5-1   
 Selected Mineral and Vitamin Comparison (mg/100g edible portion)
 83 of Iva annua to Two Modern Asteraceae Seed Crops
Table 5-2
 Collection Locations and Dates 87Table 5-3
 Parental Heritage of the C:N Ratio Samples
 88Table 5-4
 The Origin of Samples for GC-FID Analysis
 90Table 5-5
 Mass Percent of Amino Acids
 93Table 5-6
 Iva annua kernel quantities required to meet USDA minimum
 103 levels of essential amino acids                 ixLIST OF FIGURES
Figure 1-1 
 Iva annua 
 3Figure 1-2 
Iva annua
Figure 1-3
 Iva annua
Figure 1-4 
 Nutritive Value of North American Seed Crops of Prehistoric 
 Eastern Woodlands
 8Figure 1-5 Range of Extant Populations of 
Iva annua 9Figure 1-6
 Occurrence of archaeological 
Iva annua through time
 14 Figure 2-1  The Taxonomic History of the Genus 
Iva 32Figure 3-1
 Linnaeus™s original 
Iva species descriptions in Species Plantarum 53Figure 3-2 Jackson™s illustration (1960) of Linnaeus possible lectotype of 
Iva annua 54Figure 3-3 Lectotype of 
Iva annua L. from the Linnean Society of London
 55


Figure 3-5
 The syntype of Iva ciliata variety latifolia de Candolle 62Figure 3-6
 The Iva caudata syntype 63Figure 3-7
 Cypselae from 
Iva ciliata (now annua) var. 
macrocarpa 68Figure 3-8 
 Cypselae size spectrum from Little Salt Fork, Lancaster Co., Nebraska
 68

  
of I. annua var. 
caudataFigure 4-1 Bract Ratios of 
Iva annua Herbarium Specimens 78Figure 4-2 The range of the present varieties of 
Iva annua 79Figure 5-1 Comparison of Mass Percent of Proteogenic Amino Acids 
 93  
by Freshwater and Saline Sites
Figure 5-2 Sources of the main signals depicted in an NMR spectrograph
 95Figure 5-3
 Percent lipid in cypselae of populations from fresh and saline sites
 96xFigure 5-4
 Percent TAG vs Free Fatty Acid
 96





Figure 5-6 
 Box-plots of the 4 Iva Fatty Acids by Percent
 97Figure 5-7
 A characteristic result from the GC-FID analysis process
 98Figure 5-8
 Accumulated Percent Germination at Three Salinities
 99Figure 5-9 Component Fatty Acids in Major Asteraceae Oil-Seed Crops
 105Figure 5-10 Principal component analysis of the fatty acids
 108Figure 5-11  Percent Lipid by Cypselae Weights
 109Figure 5-12  Percent Nitrogen by Cypselae Weights
 1091CHAPTER 1The Biology and History of Marshelder, 
Iva annuaIntroduction
In 1924, in a preliminary account of an archaeological expedition to the Ozark region in the north-west corner of Arkansas, by M. R. Harrington  (Harrington 1924), he acknowledged that there were 
several places in the United States, east of the Mississippi, namely in Kentucky and Tennessee that 
had caves or dry rock shelters where the dryness of the sites have preserved items usually lost to the that obtained by our Ozark expedition,fl (Harrington 1924, page 1).From this introduction Harrington later announced the discovery, that along with the large invento
-restfl (Harrington 1924, page 6). Beginning with this intriguing note in 1924, knowledge of a newly discovered food and crop plant species began to emerge.  The bulk of these 
 seeds turned out to belong to the species we now call Iva annua; in the world of common names; marshelder, 
sumpweed, seacoast marshelder, and rough marshelder. 
DescriptionIva annua, is an annual (see formal taxonomic description in Chapter 2 Taxonomy), reaching 
between 0.6 Π2.0 meters (2 Π6 feet) in height. There are 
Iva annua populations that are restricted to 
2tolerable salinity 0.5-0.7 percent NaCl (Ungar and Hogan 1970). Those plants seem to reach a height 
from about 0.36 to 1 meter (Kaul 2006 and personal observation). Plants growing at the margins of a 
freshwater environment reach the uniformly taller stature of 1.25 to 2 meters (Rydberg 1922).

described as ‚hispidulous™ (Kaul 2006), because of the covering of hairs on many plants. These hairs are 
usually white and vary in length from 0.5 mm on some plants to 3.0 mm in length on others (personal 

loss of vigor late in the season, these trichomes become stiff and glassy and capable of penetrating the skin 
of the hand as an irritant (personal observation). This is almost certainly what Diamond (1999, page 151) 
meant when he said it ficaused skin irritation.flThe three-veined leaf is rounded or tapered at the base, with the distal end terminating in an acuminate, 
but very sharp pointed tip. The leaf margins are articulated into shallow, variable, rather rounded teeth, 
that may in the extreme, be virtually absent on a particular leaf (Kaul 2006, and personal observation). 

-tending™ bracts. The involucre is somewhat hemispheric composed of 3-5 phyllaries. The earliest pistillate 

-
but within the involucre (Figure 1-1). Although Jackson (1960) described the 
Iva annua 3causing an anomaly of this nature (see also Figure 3-4). In live collections from Illinois, Kentucky comprised of the ovary and the style, ending in these two elongate stigmatic extensions. During an-thesis, it is only these elongate pistil lobes that extend beyond the involucre. These are usually white 
reinforcing the place where the two pollen bags met are the only remaining structures after pollen Figure 1-1
 Iva annua
 in˜orescence and capitulum 
A
-
B- Anatomy of the capitulum of the 
Iva annua 

AB4early November. Some plants, that may have been producing moderate amounts of reddish or purple 
-ing either partially or almost completely deep reddish purple to dark purple, approaching what passes for ‚black™ on a plant (personal observation).
The mature capitula with seeds are often still on the plant after it turns brown. In the capitula from which I collected cypselae at Wickliffe, Kentucky, on a site adjoining the Mississippi, there was a 
large tendency to shatter (easily break away from the plant and disperse the fruits) compared with 
ones I collected further north near Granite City, Illinois. 
Archaeology
We now know that this species was used as a food seed, nurtured amongst ancient American Indians 

Figure 1-2 
 Iva annua 
pistillate ˜oret devel
-opment 


Figure 1-3
  Iva annua 
staminate ˜oret develop
-ment 
pollen dispersed and the connectives desiccated. 5skills to produce vastly larger seeds than their wild antecedents, and was ultimately lost in their lexicon 
of food resources as a crop, and even as food plant. In North American Ethnobotany, Moerman (1999) 
lists American Indian uses for 
Iva axillaris as a source for several medicinal uses, mostly along the 
lines of dermatological treatments, birth control, and some digestive applications; and Iva xanthifolia as 


Iva 1931, Melvin Gilmore, Curator of Ethnology at the Museum of Anthropology at the University of 
Iva xanthifolia (Gilmore 1931). One of the problems in determining these materials, was that no one up to this time had seen seeds, cypselae from Iva that were approximately 400 percent larger than wild populations. Most of the literature calls the fruits of 
ovary (although both terms are considered correct). Gilmore later reassessed these cypselae as being from Iva ciliata Willd.
, now regarded as a junior synonym of Iva annua L.In Gilmore™s description of the fiunknown seedsfl found in the caches in Arkansas (Gilmore 1931, 
page 101), he adds, fiA very interesting and curious fact is that the seeds in the stores were of a size 
much larger than any now growing as weeds.fl This suggests that all these larger seeds in the stores 
of the Bluff-Dwellers may have been the product of cultivation. The purpose for which they were 
used is problematic.fl In a letter to Gilmore, Safford (1924) postulated that perhaps 
Iva annua seeds were fitoo acidulous and astringent for foodfl and suggested that this species could have provided 
Anthropology at the University of Michigan could not locate Dr. Safford™s letter (Letter to M. R. 
6Gilmore; Concerning Specimens of Ozark Bluff-Dweller material collected by M. R. Harrington) to 
Gilmore on this subject.]This of course, led to a discussion about for what exact purpose these caches of ancient seed were stored. Clearly, as far as known, they had reached their maximum size during the period of use as 
crop production by these indigenous peoples, as demonstrated by archaeological discovery. 
However, Volney H. Jones, mentioned (Jones 1936) that identical large seed remains had been 
found at the Newt Cash Hollow in Menifee County, Kentucky. He writes, fiThese seeds occur spar
-ingly throughout the material but comprise a rather high percentage of the fecal matter,fl (Jones 1936, 
were ingested without removing the seed coat. By the time that Asch and Asch (1978) produced their 
study of I. annua nutritional components, this debate was fairly settled. Later archaeologists discov-ered the presence of human feces as a by-product of activities during the Woodland period, within 
the great midcontinent cave systems of the Midwest (Faulkner 1991, Gremillion 1996, Watson and 
Yarnell 1966)  The fact that sumpweed seeds comprised a large percentage of the volume of these 
feces underscored the importance of I. annua as food in these societies. When Asch and Asch (1978) 
published their study on the economics of using sumpweed as a food seed, including an analysis of its high levels of protein and oil, the compelling case for Iva annua as a food plant was well ac-cepted. I. annua is a high oil, high protein food seed (Asch and Asch 1978, and Chapter 5 here) that has 
been found in association with humans in Eastern North America for some 7000 years (Wagner and 
Carrington 2014). It shows signs of having been cultivated for about 4500 years. Unlike most do-7mesticated food plants, by the beginning of the 20th century, it had fallen out of the repertoire of not 
only crop plants, but had disappeared from the list of edible plants used as American Indian foods 
in North America. There are few other exxamples (except for possible varieties in the grass fam
-ily (Poaceae), and perhaps the Iva relative giant ragweed, 
), where its food use was discovered solely through archaeological discovery. None of the chronicles of American Indian food 
plants, and indeed neither the compendia entitled Sturtevant™s Notes on Edible Plants (Sturtevant 
1919), nor Tanaka™s Cyclopedia of Edible Plants (Tanaka 1976) nor any other published collection of 
Iva annua as a food resource. The sites where I. annua cypselae are found in a archaeological context also tend to include evidence of a number of other plant species believed to have contributed to the subsistence of the prehistoric indigenous people who inhabited these sites. This group of plants is known as the Eastern 
Agricultural Complex. The core group of the Eastern Agricultural Complex (EAC), are six species 
of food seed grown starting from approzimately 4500 to 3000 BC, although not necessarily by all groups practicing cultivation, and certainly not uniformly as the same list of plants (Smith 2006, Yarnell 1993). They are a goosefoot annual, 
a smartweed, erectum, 
a maygrass, 
a barley, 
Hordeum pusillum, 
a marshelder, 
Iva annua, 
 Helianthus annuus. This suite of species also includes a cucurbit, (McConaughy 2008). Figure 1-4 shows the approximate nutritional components of  the six important seed foods (excluding ) listed above compared to Zea mays.In addition to these plant species, there are up to 13 other species in various levels of consideration 
to be considered members of the EAC (Yarnell 1993). Some of these candidates include: 

, Strphostyles helvola, , 
, Apios americanus, -8
, Solanum nigrum (complex?), 
, Mollugo verticillata, maculata, plus others. Of those just mentioned, certainly could be the most problematic as it is a well-known toxic plant for people and animals (Kingsbury 1964, Wagstaff 
2008). Although since it shares habit and habitat with 
and sometimes literally overlap on adjacent locations (personal observation), accidental contamination seems like a possibil-ity (personal opinion). In today™s world, 
Iva annua is considered a weed, and is not terribly well-known outside areas of the American South and adjoining Great Plains, Figure 1-5. It tends to be found in disturbed ground 
regimes especially along water ways, river terraces, and stream bottomlands. The domesticated 
0102030405060708090100LipidProteinCarbohydrateFiberAshIva annua [Asch & Asch 1978]Chenopodium berlandieri [Asch & Asch 1985]Helianthus annuus [Watt & Merrill 1963]Hordeum vulgare [Duke & Atchley 1986] (No available analysis for Hordeum pusillum)Phalaris caroliniana [Crites & Terry 1983]Polygonum erectum [Asch & Asch 1985]Zea mays [Asch & Asch 1978] Nutritive Value of North American Seed Crops of Prehistoric Eastern WoodlandsFigure 1-4 
 Nutritive Value of North American Seed Crops of Prehistoric Eastern Woodlands 

Zea mays. 
Hordeum vulgare 
Hordeum pusillum was located.9Walsh Co., NDBeadle Co., SDDavison Co., SDOakland Co., MIKent Co., MIIva annua var. caudataSocorro Co., NMIva annua var. annuaCarrington™s cypselae sourced from: (lf. to rt.)Lincoln, NebraskaGranite City, IllinoisWickli˜e, KentuckyBloomington, IndianaColumbia, South CarolinaFigure 1-5 
 Range of Extant Populations of 
Iva annua
 -tion of Iva annua 
The location in New Mexico, at the Bosque del Apache Wildlife Refuge is a sigle record from the 


railroad yard.
10strains of Iva annua are presumed to be extinct (Blake 1939, Wagner and Carrington 2014). But the 
existence of Iva annua kernels in the archaeological record is very rich. Twenty years ago it was pre
-sumed that the cultivation of domesticated Iva annua had died out sometime around the year 1100-
1400 AD, but more recently there have been recovered charred cypselae and kernels from sites dated 
from the Middle Archaic (cal. 5970-4945 BC) to the period of contact in the late AD 1700s, and per
-haps into the 1800s (Wagner and Carrington 2014). The geographically extent of these discoveries 
extends from southern Ontario and central North Carolina, along the Gulf Coast to Southern Missis-sippi, to the panhandle of Oklahoma and Texas and up into the Canadian Plains (Wagner and Car
-rington 2014). These often charred remains are probably comprised of seed dropped from meals and 
cleaning events and to some extent may even represent seed from some of the weedy representatives with which these groups shared the environment. Importantly, 
Iva annua seed have also been recov-ered from woven bags of cached seed, possibly being saved for planting and, most fortunately for archaeologists, from desiccated feces left in caves, especially including those in the Mammoth, Salt, and Big Bone Cave systems in Kentucky and Tennessee (Faulkner 1991). These ancient meals have 
occasionally been augmented by the discovery of mummies that also contained consumed cypselae.The unusually large cypselae that led to the exploration of 
I. annua as a cultigen, were given va-rietal status (Blake 1939), Iva ciliata var. 
macrocarpa,
 because of their striking size departure from the Iva seeds with which the modern world is familiar. When Jackson revised the genus 
Iva (Jackson 1960) this became the new combination Iva annua var. 
macrocarpa. 
Its position as a domesticate derived from its very large cypselae, 4.8 mm by 3.2 mm in size or larger, its association with well-
known domesticated plants in intentional storage contexts, and a geographical distribution in ar-11chaeological sites that was thought to extend further to the north and east than the present range for the distribution for wild I. annua (Black 1963, Blake 1939, Gilmore 1931, and Jones 1936). More I. annua than was thought (Figure 1-5). Pollen cores from the Holocene of Moon Lake, show that I. annua was common for a time as far north as North Dakota, reaching its peak between 8000 cal. yr BP and 
7000 cal. yr BP. At present, there is only one collection locality for North Dakota, and the species is 
In R. C. Jackson™s revision of 
Iva (Jackson 1960) he includes an additional variety, 
Iva annua var. 
caudata. I demonstrate in chapter four, The Taxonomic History of 
Iva annua, and the synonymy of Iva annua var. 
caudata (ASTERACEAE), that this variety should be considered synonymous with Iva annua var. 
annua; therefore this variety does not appear as a separate entity on the range map for modern Iva annua shown in Figure 1-5.Iva Cypselae Over Time
The archaeological record of marshelder use extends back to at least 4500 years before the pres-ent (Wagner and Carrington 2014). The maps (Figures 1-6 A-F) show the geographic range of its 
use from the oldest archaeological discoveries through the protohistoric-historic sites that mark the end, so far, of its most recent presence in the cultural sites of eastern Native Americans. Although the 
classical designation for the time that I. annua disappears from the record is around 1000 AD (Smith 
1989), more recently it has been established that I. annua was cultivated in a few places to around 1820 (Wagner and Carrington 2014). 

12and that they had nurtured for over three and a half millennia. The obvious possibility is that when 
maize and the crops from the Southwest came, they dropped the older suite of plants for the high-
I. annua was cultivated into far later dates than were originally surmised (Wagner and Carrington 2014) we are 
also learning that maize use in the same region predated our earlier notions.Archaeological maize research was done by searching for the macro-remains of mostly cobs for decades. What has so increased our perspective on maize over the centuries of its use is the more 
recent understanding of its microfossil remains; namely pollen, phytoliths, and starch granules (Hart et al. 2007). Pollen evidence clearly implies that a crop was growing nearby since pollen is produced by the maize plant weeks before it ripens edible grain. Phytoliths and starch grains however, are 
recovered from residues in vessels ostensibly used for food preparation. Starch grains are clear evi-dence of cooked corn kernels, but phytoliths come from soft glumes or chaff, cob features, (Pearsall 
et al. 2004) and are not irrefutable proof of maize grown on-site. Although one would not waste the 

from the cobs, either for cooking or for transport would both result in deposits of phytoliths when in the cooking pot for later analysis. Maize phytolith evidence has moved claims of maize cultivation back as far as 2270 B.P. in Central New York State (Hart 
et al. 2007), and as far back as 1515 B. P. 

previous notion of maize culture becoming a major contribution only from approximately 1000 to 1150 A.D. (Smith 1992). But then Boyd and Surette (2010) claim on the basis of phytolith evidence 
that maize was in more or less continuous use in the Canadian boreal forest from 300 B.C. There 
13could be an issue here, especially involving using phytolith evidence to support a claim for maize cultivation in the Canadian boreal forest. Today, most of the range of the boreal forest is above the 

a regular part of the ‚boreal™ diet?  Clearly, Native American trading networks were widespread and 
robust (Turner and Loewen 1998). When one thinks of Indigenous people trading maize, some tend 
to imagine a person carrying a small bag of maize. But when one reads of Anasazi hand-carrying 
thousands of 600 to 800 pound logs for journeys of ten days or more (Betancourt et al. 1986), one incentives are right. My view is that since the macro-remains have not been found to support the adundant micro-remains, claims of maize culture, as aginst use, lack enough credible evidence. If we take macro-remains of maize culture, strongly supported by cobs, or even pollen, as the basis for assigning a probable time span for the origin of maize culture, then the spread of maize farming is more credible for gradually changing the pressure to keep or abandon Iva farming as we consider dates moving forward from 1000 A.D..
Measuring Iva annua CypselaeIn order to trace the ever changing sizes of Iva annua fruits through the endeavor of archaeology, it 

measurements.  A large number of the recovered 
I. annua cypselae that have been obtained from the relevant sites were charred. So it is necessary to know how paleoethnobotanists have made size determinations for these objects. Experiments on the kernels and whole cypselae of I. annua have 14



F Early Mid Miss./Mid Ceramic
Figure 1-6  Occurrence of archaeological 
Iva annua 
through time
A-H depict the locations of I. annua cyselae at archaeological sites in the eastern United States and 
adjoining Canada from the 
 Middle Archaic through the Historic.

15



shown that when they are charred or carbonized, they shrink (Asch and Asch 1985, Wright 2008, 
Yarnell 1972). These workers have performed experiments on both kernels (the meat of the seed 
itself) and whole cypselae (the entire fruit including the kernel and the pericarp; in conversation the 
Hopefully to yield predictable enough results to standardized a correction factor that could be used to more accurately compare the measurements of cypselae as the sizes changed through the centuries under cultivation. This correction factor is also sought to aid in comparison of materials that have 
been desiccated with those materials that were carbonized by heat. In Blake™s original description 
of var. 
macrocarpa 
were diagnostic of the varietal designation, but today paleoethnobotanists recognize cypselae lengths (including reconstructed) of 4.0-4.2 mm as the presumed lower limit of cypselae size in domesticated I. annua (Asch and Asch 1985, 161-162, Smith 1987, 1992, 49).

-Figure 1-6 (Cont™d)16ized kernels are smaller than carbonized cypselae by 0.7 mm in length, and by 0.4 mm in width, and that 
carbonized cypselae are smaller than non-carbonized cypselae by 10 percent in both length and width.Later, and in order to somewhat standardize the comparisons between 
I. annua
Helianthus annuus, Yarnell (1978) used the size index, the product of the reconstructed length times 
width. This meant that in the example of wild/weedy 
I. annua cypselae from an east-central Missouri site that produced cypselae with a mean length (2.9 mm) times width (2.5 mm) yielding a size index of 

the specimen condition, Yarnell (1978) found the size indices from the terminal Late Archaic (indices 
= 8-12), during the Early Woodland  (indices = 12-16), Middle Woodland (indices = 16-20), the early 
Late Woodland (indices = 20-26), and during the Mississippian Period (indices = 25-40). This works 
out to approximately 1 mm of cypsela length increase per 1000 years (Asch and Asch 1978, page 323).
Additional charring experiments by Asch and Asch (1978, 326) indicated that the larger the re
-covered kernel, the larger the correction factor needed to simulate the original dimensions. With this 

of the archaeological cypselae. The correction factor that they (Asch and Asch 1978) calculated for 
estimating the uncarbonized dimensions from the charred specimens is thus: Cypsela length uncarbonized = 1.36 x (kernel length carbonized) + 0.17 mm, Cypsela width uncarbonized = 1.45 x (kernel length carbonized) - 0.06 mm.
yield a calculation of size indices noticeably greater than that obtained using Yarnell™s factor (e.g. N. 
17
formula, and some workers (Adair pers.comm. with Gail Wagner, 2010) still compensate for shrink
-age when they multiply the charred cypsela length by 1.11, and by whatever reckoning, these tech
-
Patti J. Wright™s experiments with 
I. annua and H. annuus cypselae being roasted and carbonized under controlled conditions (Wright 2008) and over a range of temperatures and exposure times has 
provided a high-resolution picture of the relationship between temperature, treatment times, anaero-bic conditions and mass loss and shrinkage. Her detailed study produced from over 1400 I. annua cypselae, heated at a selection of temperature intervals has greatly improved our understanding of what heating regimes resulted in which changes in cypselae dimensions. Wright also points out 

render the materials so friable as to greatly reduce the probability of surviving the mechanical dam-age accompanying burial in an archaeological context. She goes on to point out this indicates a very -tion; hence the small number of I. annua and H. annuus seed preserved when compared with those of starchy seeds that are represented at these hearths in a more robust fashion.Wright™s (2008) contribution to the correction standards discussion recognized that although Yar
-nell™s corrections factors underestimated the size of 
I. annua cypselae somewhat, a point he himself admitted, the contribution of these considerations still helped establish the recognition that cypselae 18were increasing in size throughout the time that I. annua was under cultivation in the North Ameri
-can midcontinent, and that this principle still is recognized to be true. Wright™s recommendation that 
an upgraded range of compensation factors; 1.08-1.30 percent for correcting length, and 1.13-1.47 percent for correcting the widths of charred kernels to estimate the original size is consistent with the results of her extensive experiments. Although cypselae of a reconstructed length of 4.0-4.2 mm 
(Smith 1987, page 23), and the cypselae of modern wild I. annua have been reported to range up to 5.4 mm in length (Asch and Asch 1978, page 322). It is reliable to assume that an assemblage of 
I. annua cypselae is of domesticated origin, only when the small end of the size distribution of the population is near 4.0 mm in length.Minerals and Vitamins
In addition, Asch and Asch (1978) provided additional nutritional data for 
Iva annua with the vitamin and mineral components listed in Table 1-1 and compared 
I. annua to two common oil seed plants also from the Asteraceae, 
Helianthus annuus L. and Carthamus tinctoriusgeneral, the nutrient content of I. annuatwo other oil seed plants.  Asch and Asch (1978) reported 
I. annua seeds contain approximately one third (29%) of the Recommended Daily Intake (RDI) for a day™s worth of calcium consumption (per 
100g). The USDA National Nutrient Database for Standard reference Release 27 puts broccoli, 
Bras-sica oleracea
edible portion as mg/100 mg EP). Broccoli is considered an excellent source of calcium, even at that 19level. Table 1-1 reports levels of phosphorus and potassium from 
Iva annua at very high levels, but limiting factor of most human diets. The amount of iron contained in I. annua seed was reported by Asch and Asch (1978) to be 11.4 
mg/100 mg EP providing just under 2/3 of the RDI. This sounds like an elegant solution to the chal
-provide easily available (absorbable) iron in most cases, it does not simply follow that the majority of the iron in an analysis could be biologically available (Hurrell and Egli 2010). 

Thiamin is considered safe at relatively high levels and in fact there is no tolerable upper intake 
Asch (1978) reported that in a 100g serving, the kernels of Iva annua would provide approximately 
 Selected Mineral and Vitamin Content (mg/100g edible portion) of Three 
Asteraceae Seeds Grown as Crops
Iva annua, Marshelder*CalciumPhosphorusIron%DV%DV%DV%DV%DV%DV%DVPotassiumThiaminRibo˜avinNiacinVitamin B1Vitamin B2Vitamin B3290130011.47802.130.7513.1Helianthus annuus, Sun˜ower*1208377.19201.960.235.4Carthamus tinctorius, Sa˚ower77.86642.865685.711000mg1000mg18mg3500mg1.0710.3572.1431.5mg1.7mg20mgSelected Mineral and Vitamin Content (mg/100g edible portion) of Three Asteraceae Seed Crops* Numbers taken from:  Asch and Asch, 1978 (Table 3, page 307) Numbers taken from:  nutritiondataself.com/facts/nut-and-seed-products/3068/229%12%7.79%130%83.7%64.3%63.3%39.4%27.8%22.3%26.3%19.6%142%130.7%71.4%44.1%13.5%21%65.5%27%10.7%Recommended Daily Values˛ (www.dsld.nlm.nih.gov/dsld/dailyvalue.jsp)˛ Numbers taken from: www.dsld.nlm.nih.gov/dsld/dailyvalue.jspIva annua, Marshelder*CalciumPhosphorusIron%DV%DV%DV%DV%DV%DV%DVPotassiumThiaminRibo˜avinNiacinVitamin B1Vitamin B2Vitamin B3290130011.47802.130.7513.1Helianthus annuus, Sun˜ower*1208377.19201.960.235.4Carthamus tinctorius, Sa˚ower77.86642.865685.711000mg1000mg18mg3500mg1.0710.3572.1431.5mg1.7mg20mgSelected Mineral and Vitamin Content (mg/100g edible portion) of Three Asteraceae Seed Crops* Numbers taken from:  Asch and Asch, 1978 (Table 3, page 307) Numbers taken from:  nutritiondataself.com/facts/nut-and-seed-products/3068/229%12%7.79%130%83.7%64.3%63.3%39.4%27.8%22.3%26.3%19.6%142%130.7%71.4%44.1%13.5%21%65.5%27%10.7%Recommended Daily Values˛ (www.dsld.nlm.nih.gov/dsld/dailyvalue.jsp)˛ Numbers taken from: www.dsld.nlm.nih.gov/dsld/dailyvalue.jsp20
survival resource with respect to this essential nutrient.can cause such conditions as sore throat, edema of the pharyngeal and oral mucus membranes, and 
Reference Intakes 1998). Niacin, vitamin B3, is primarily a coenzyme for transferring hydride ions in the presence of several 
dehydrogenases. The main consideration in establishing the recommended dietary allowance (RDA) 


niacin, in addition to the direct dietary intake values. The RDA for adults is 16mg/day of NEs for men 

I. annua 
kernels at somewhere between 80 and 94 percent, depending upon gender of the adult. The tolerable 
upper limit per day of niacin is set at 35 mg/day for an adult (Institute of Medicine (US) Standing Com-
(facial reddening), not to be ignored, but as much a cosmetic as potentially lethal reaction.
All in all, these measurements make Iva annua kernels among the most well-provisioned for the B-vitamins and calcium of known seed foods. But, as impressive as these measurements are in this realm, it is the yields in protein and oil that have commanded the most attention (see Chapter 5).21Research Question Number 1.
 When the nutritional components of the seeds of 
I. annua, as as-
areas that should be examined against the needs of modern agriculture? The initial analysis (Asch 
and Asch 1978) has been cited numerous times in introducing discussions about ancient crop species. 
Research Question Number 2.
 Will adding to the picture of 
I. annua as a protein and oil plant be 

behaviors in any substantive way in our understanding of the dynamics of the EAC communities?Research Question Number 3.
 The designation of three varieties of 
I. annua, seemed arbitrary and incomplete. Especially for the two extant varieties, I. a. annua and I. a. caudata, does a careful morphological examination of these varieties bear out the taxonomy behind their establishment, or should these designations be recognized as a synonymy, thus ending the discussion started by Small 
(1899)  by the description of I. caudata?Research Question Number 4.
 This species, occurs in both saline and freshwater environments. 
Do the populations in these different habitats have differences in their nutrient production that 
represent a resource that could be tapped in the search for food production in areas that suffer salt-
contaminated soils? Chapter 2 describes the taxonomic history of the genus Iva. Since one of the drives for the botani-cal exploration of the New World was the discovery of new medicinal plants, 
Iva with its camphor smell was on the ‚radar™ early for European physicians, even before Linnaeus.
Chapter 3 provides the taxonomic history of the species Iva annua itself. Although it was one of 
22Iva, mistaken location data and misinter-pretation led to confusion and synonymy that took two centuries to be fully resolved. Buried within the case-study of this small, not-very-well-known genus is a microcosm of many of the issues that debates of today. This includes a synopsis of the history of 
Iva annua from the years before the pub-lication of in binomial nomenclature, and includes many of the greatest botanists in the history of eighteenth century Europe.Chapter 4 is a detailed look into the details of the modern taxonomy of Iva annua. This is where the discussion of Research Question number 3 is carried out. I present the investigation into the controversial species that later became a controversial variety. Here is presented a history of what I 
claim by chapter™s end, is the synonymy of one of the extant subspecies in Jackson™s reorganization 
(1960) of the genus Iva. Chapter 5 provides some new insight into the possible nutritional and physiological traits of this paper (Asch and Asch 1978) has provided virtually all of the nutritional information that is cited 
about this species and its vitally important role in the diets of the cultures that developed this species from a weedy annual found along streams, rivers and coastlines, into a highly nutritious adjunct to the diet and food supply system of the prehistoric Native Americans east of the Mississippi river. In 
this critical publication (Asch and Asch 1978), only one sample of 
Iva annua cypselae was analyzed to produce the results that are so well-known today. Chapter 5 updates this information and adds the 
23of this food seed. In a series of germination studies, I start to unravel the roles of salt tolerance in those populations that are concentrated in the areas of the prairie West where this species is primarily 
an inhabitant of the medium salinity zone (Ungar and Hogan 1970) of the saline wetlands of Kansas and Nebraska and other points in central North America.
24REFERENCES25REFERENCES ASCH, D. L., A
SCH, NANCY B. (1985). fiPrehistoric Plant Cultivation in West-Central Illinois.fl 
  
 University of Michigan 75: 149-203. (1978). fiThe economic potential of Iva annua and its prehistoric importance in the lower  Illinois Valley.fl 

, University of Michigan   67: 300-341.ASCH SIDELL, N. (2008). The Impact of Maize-based Agriculture on Prehistoric Plant Communities 
  in the Northeast. 
 J. P. Hart. Albany, New York, New 
  York State Museum. 512: 29-51.
BETANCOURT, J. L., DEAN, JEFFREY S., HULL, HEBERT M. (1986). fiPrehistoric Long-Distance  Transport of Construction Beams, Chaco Canyon, New Mexico.fl 
 American Antiquity 51(2): 370-375.BLACK, M. (196 Iva annua L.fl 
 48: 541-547.BLAKE, S. F. (1939). fiA New Variety of Iva Ciliata From Indian Rock Shelters in the 
 South-Central United States.fl 
Rhodora 41(483): 81-86.BOYD, M., SURETTE, CLARENCE (2010). fiNorthernmost Precontact Maize in North America.fl 
 American Antiquity 75(1): 117-133.
DIAMOND, J. (1999). Apples or Indians. 
Guns, Germs, and Steel. New York, W. W. Norton and 
  
  Company: 131-156.
DUKE, J. A., A
TCHLEY, A
LAN A.
 (1986). 
.   Boca Raton, Florida, CRC Press, Incorporated.FAULKNER, C. T. (1991). fiPrehistoric Diet and Parasitic Infection in Tennessee: Evidence from the 
  Analysis of Dessicated Human Paleofeces.fl American Antiquity 56(4): 687-700.GILMORE, M. R. (1931). fiVegetal remains of the Ozark Bluff-Dweller culture.fl 
Michigan Academy 
 
14: 83-102.GREMILLION, K. J. (1996). fiEarly Agricultural Diet in Eastern North America: Evidence from Two 
  Kentucky Rockshelters.fl American Antiquity 61(3): 520-536. GRIMM, E. C. (2001). fiTrends and Palaeoecological Problems in the Vegetation and Climate 
 History of the Northern Great Plains.fl Biology and Environment: 

 Royal Irish Academy 
101B (NO. 1-2): 47-64.26HARRINGTO
N, M. R. (1924). fiThe Ozark Bluff-Dwellers.fl 
American Anthropologist 
26: 1-21.HART, J. P., B
RUMBACH, HETTY J., LUSTECK, ROBERT (2007). fiExtending the Phytolith Evidence  for   Early Maize (Zea mays ssp. mays.) in Central New York.fl 
 American Antiquity 73(3): 563-583.HURRELL, R., EGLI, INES (2010). fiIron bioavailability and dietary reference values.fl The American  91(suppl): 1461S-1467S.INSTITUTE OF MEDICINE (US) STANDING COMMITTEE ON THE SCIENTIFIC EVALUATION OF DIETARY  REFERENCE INTAKES AND ITS PANEL ON FOLATE, O. B. V., AND CHOLINE (1998). 

 Washington, D.C., National Academies Press.
JACKSON, R. C. (1960). fiA revision of the genus Iva L.fl 
University of Kansas Science  Bulletin 41: 793-876.JONES, V. H. (1936). fiThe Vegetal Remains of Newt Kash Hollow Shelter.fl 
Reports in Archaeology 
  and Anthropology
, University of Kentucky 3(4): 147-167.KAUL, R. B., SUTHERLAND, DAVID M., ROLFSMEIER, STEVEN B. (2006).   Lincoln, Nebraska, University of Nebraska.
KINGSBURY, J. M. (1964). . Englewood Cliffs, 
  NJ, Prentice-Hall, IncMCCONAUGHY, M. A. (2008). 
Current Issues in Paleoethnobotanical Research from Pennsylvania 
  and Vicinity.

J. P. Hart. Albany, The New York State 
  Education Department. II: 9-28.MOERMAN, D. E. (1999). 
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PEARSALL, D. M., CHANDLER-EZELL, K., ZEIDLER, JAMES A
. (2004). fiMaize in ancient Ecuador:   
 results of residue analysis of stone tools from the Real Alto site.fl 
 
e 31(2004): 423-442.RYDBERG, P. A.
 (1922). 
 New York, New York Botanical 
  
 Garden. 33: 3-13.SAFFORD, W. E. 
(1924). Letter to M. R. Gilmore, April 17, 1924, concerning specimens of Ozark 
  Bluff-Dweller material collected by M. R. Harrington. M. R. Gilmore. Ann Arbor, Michigan, 
  Unpublished.SALVADOR, R. J. (1997). Maize. The Encyclopedia of Mexico: History Culture and Society,
  Fitzroy Dearborn Publishers.
27SMITH, B. D. (2006). Prehistoric Plant Husbandry in Eastern North America. 

P. J. W. C. Wesley Cowan. Tuscaloosa, University 
  of Alabama Press: 101-119.
SMITH, B. D.(1992). Rivers of Change. Washington, Smithsonian Institution Press.
 (1989). fiOrigins of Agriculture in Eastern North America.fl 
Science 246: 1566-1571. (1987). The Independent Domestication of Indigenous Seed-Bearing Plants in 
 Eastern North America. 
Emergent Horticultural Economies of the Eastern Woodlands.
  W. F. Keegan. Carbondale, IL., Southern Illinois University. 1: 3-47.
STURTEVANT, E. L. (1919). 
 Albany, N. Y., State of New York, 
  Department of Agriculture.
TANAKA, T. (1976).
. Tokyo, Yugaku-sha.
TURNER, N. J., LOEWEN, DAWN
 C. (1998). fiThe Original fiFree Tradefl: Exchange of Botanical 
  
 Products and Associated Plant Knowledge in Northwestern North America.fl 
 Anthropologica 
40(1): 49-70.WAGNER, G. E., CARRINGTO
N, PETER H. (2014). Sumpweed or Marshelder (Iva annua). New Lives   for Ancient and Extinct Crops.
 P. Minnis. Tucson, University of Arizona Press. 1: 65-101.
WAGSTAFF, J. D. (2008). International Poisonous Plants Checklist. New York, CRC Press.
WATSON, P. J., 
YARNELL, RICHARD A
. (1966). fiArchaeological and Paleoethnobotanical  Investigations in Salts Cave, Mammoth Cave National Park, Kentucky.fl 
American Antiquity   31(6): 842-849.WRIGHT, P. J. (2008).
 Sumpweed Remains.fl 
33(2): 139-153.YARNELL, R. A. (1

 
, University of Michigan 67: 289-299. (1993). 
.   
 Foraging and Farming in the Eastern Woodlands. C. M. Scarry. Gainesville, 
 University of Florida: 13-26.
 (1972). fiIva annua var. 
macrocarpa
: Extinct American Cultigen?fl 
 American Anthropologist 
74: 335-341.28CHAPTER 2A Short Taxonomic History of Genus 
Iva (ASTERACEAE)Iva were assigned to it came from the eighteenth century work of Carl Linnaeus. The rules of modern 
taxonomy, provisionally set at the International Botanical Congress of Vienna in 1905, literally 
dictate the beginnings of plant taxonomy as starting with Linnaeus™s 1753 publication of 
Species Only names established in this work, or afterwards according to its example are taken as legitimate. Notwithstanding this convention, many plants, including many from outside Europe were already well known and had a history in the European consciousness predating the publication of  (Linnaeus  1753).In the eighteenth century, as many regions came under the preview of European exploration, it was 
plants, especially those of possible medicinal importance that commanded a share of the attention during these explorations that was also shared with discoveries of precious metals and agricultural regions.Iva annua, was in the 1719 publication by Sébastien Vaillant (1719), titled 


, under the name Tarchonanthos 
(Vaillant 
1719, page 310). Here Vaillant uses the name 
Tarchonanthos. 
modern genus name itself in current literature have this name spelled Tarchonanthus.
Sébastien Vaillant was a surgeon in Paris who studied botany under Joseph Pitton de Tournefort 
29and later made important contributions to botany. He lists two species of 
Tarchonanthos, 
one from America, and one from Africa; he also referred to them as 
Conyza Americana and Conyza Africana, 
-tions in his former teacher™s work, 

(Tournefort 1700, page 455)
. Vaillant 
described the African Tarchonanthos as odorous. Today this plant is believed to be 
Tarchonanthus 
camphoratus, the camphor bush, a medicinal plant used extensively from Saudi Arabia to South Af
-rica because of its antimicrobial volatile essential oil (Matasyoh et al. 2007). For many people, fresh Iva annua has a similar smell of camphor, at a fainter level.
I have found no evidence of how Linnaeus received his specimens of Iva. Near the end of his succinct description of Iva annua, he seems to credit his materials to D. B. Jussieu (followed by the symbol for the sun that he used to signify the species was an annual). D. B. Jussieu™s brother Barnard 
worked extensively in South America, living for years in Peru, but not leaving any direct record of 
having collected Iva during his journey. D. B. Jussieu is known to have shared correspondence and specimens with Linnaeus on several occasions (Jarvis 2007, page 214).1753  In the Linnaeus assigned two species to this genus; Iva annua and Iva frutescens. The locality information noted with Iva annua reads, fi America meridionali,fl 
(see page 44). The same information for 
Iva frutescens read, fiVirginia, Peru.fl As of today, 
there are no species of Iva known from anywhere outside North America and the Caribbean. The er
-roneous locality datum for 
Iva annua led to confusion and synonymy that would not be resolved until 
the Revision of the Genus Iva by R. C. Jackson (1960). It is my opinion that the locality information for Iva frutescens, namely Virginia (as well as Peru) gave enough true footing to the species as to not 
30have presented the stumbling block as did the error in the Iva annua description. Much of the remainder of this discussion of Iva taxonomy will reference the diagram (Figure 2-1) in which I have graphically summarized the key events in the unfolding taxonomy of this genus. 1788 The next additions to the genus 
Iva were the descriptions in 1788 by Thomas Walter (1788) 
in Flora Caroliniana of Iva monophylla and The species is still recognized as a valid species of Iva, but I. monophylla was soon recognized as belonging in the genus 
 (ragweeds) and was never included within the genus Iva 
similar to 
ones, while Iva-onym of 
in 1836 (DeCandolle 1836). Today this taxon is known as 

artemisiifolia L., annual or common ragweed.1804  The next major milestone is the publication of 
Iva ciliata by Willdenow (1804). This rede
-
Iva annua in South America (See the Taxonomic History of 
Iva annua page 44). This name would stand until the reorga
-nization of genus Iva (Jackson 1960).1814  Frederick Pursh in his Flora Americae Septentrionalis
 describes Iva axillaris, in the Sup-plementum at the end of the publication (Pursh 1814, page 743). In his mention of (page 580) seems to reference a species called Iva integrifolia in a manuscript by Banks in the same paragraph. There does not seem to be any formal description for 
I. integrifolia, and it disappears until formally synonymized with in Jackson™s revision (Jackson 1960, page 815).
1818  Nuttall in his 
 (Nuttall 1818) describes Iva xanthifolia. It 31becomes one of the two most ‚synonymized™ species in genus 
Iva. Curiously, when it appears, 66 
years later, in Asa Gray™s Synoptical Flora of North America, it is spelled 
fixanthiifolia.fl This spell-ing is randomly switched with Nuttall™s original spelling until Jackson™s revision (Jackson 1960) 
when the original I. xanthifolia (with one ‚i™) becomes the usual spelling again.1820  Humboldt, Bonpland, and Knuth described I. cheiranthifolia from near Havana in Cuba (Humboldt, Bonpland, and Knuth 1820). They published plentifully, and were usually abbreviated 

simply labeled fiFloridafl that comprised the only evidence that Iva cheiranthifolia was found in the United States. Although it is listed in De Candolle™s 

-, (De Candolle  1836), (Figure 2-1, column II) it does not appear in Asa Gray™s 1884 Synoptical 
Flora of North America (Gray 1884). I have wondered if they (H. B. K.) considered the Caribbean 
outside the boundaries of North America at the time of this work.
1830  Lessing describes I. asperifolia from Veracruz, Mexico, in Linnea (Lessing 1830). Perhaps 
because Mexico was not unanimously considered to be North America, this species was ignored in 
American literature until Rydberg (1922). It has since been found growing in a single Florida county, 
but Jackson (1960, page 804) thinks it is probably introduced.1836  Augustin Pyramo De Candolle in his 
 lists the Iva known until 
his publication (Figue 2-1,column II), and adds a subspecies to the record of Iva ciliata. latifolia. De 
Candolle™s ‚wide-leaved™ variety is not exceptionally broad
. I have seen no reference (except for Jackson 
1960, page 807) to Iva ciliata . latifolia outside its 1836 appearance (De Candolle 1836).
The Prodromus of De Candolle also includes a note to include Iva angustifolia Nuttall. This name 
32Genus
 Iva
Linnaeus 1753Carl Linneaus
Linnaeus 1753Species
 annuaLinnaeus 1753Species
 frutescens
Linnaeus 1753Genus
 Iva
Linnaeus 1753Genus
 Euphrosyne
De Candolle 1836
Augustin P. de Candolle
De Candolle 1836
Species
 annuaLinnaeus 1753Species
 frutescens
Linnaeus 1753Genus
 Iva
Linnaeus 1753Genus
 Oxytenia
Nuttall 1848
Sect.
 Iva
Gray 1884
Sect.
 Iva
Gray 1884
Sect.
 Linearbractea
Jackson 1960
Asa Gray
Gray 1884
Species
 ciliataWilldenow 1804
Species
 ciliataWilldenow 1804
Species
 imbricata
Walter 1788
Species
 imbricata
Walter 1788
Species
 monophyllaWalter 1788
Species
 frutescens
Linnaeus 1753Species
 Hayesiana
Gray  1876
Sect.
 Cyclachaena
(Fresenius) Gray 1884
Species
 xanthiifolia*
Nuttall 1818
Species
 xanthifolia*
Nuttall 1818
Sect.
 Chorisiva
Gray 1884
Species 
Nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
Genus
 Oxytenia
Nuttall 1848
Genus
 Oxytenia
Nuttall 1848
Species
 acerosa
Nuttall 1848
Species
 dealbataGray 1852
Species
 ambrosiaefolia
(Gray) Gray 1884
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 ambrosiaefolia
(Gray) Gray 1884
 Genus
 Cyclachaena
Fresenius 1838
Species
 xanthifolia*
Nuttall 1818
Iva nevadensis
M. E. Jones 1883
Iva foliolosa
Nuttall 1841
Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Iva dealbata
Gray 1852
Iva H
ayesiana
Gray 1876 
Subspecies
 pubescens
Gray 1874
Subspecies
 axillaris
(Pursh 1814) Gray 1884
Subspecies
 latifolia
De Candolle 1836
Species
 axillaris
Pursh 1814
Species
 axillaris
Pursh 1814
Species
 axillaris
Pursh 1814
Species
 microcephala
Nuttall 1841
Species
 microcephala
Nuttall 1841
Species
 angustifolia
Nuttall 1836
Species
 angustifolia
Nuttall 1836
Genus
 Iva
Linnaeus 1753Per Axel Rydberg
Rydberg 1922
Species
 ciliataWilldenow 1804
Species
 imbricata
Walter 1788
Species
 caudata
Small 1899
Species
 frutescens
Linnaeus 1753Species
 Hayesiana
Gray  1876
Species
 oraria
Bartlett 1906 
Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria **
(Bartlett) Jackson 1960
  Species
 axillaris
Pursh 1814
Species
 hayesiana
Gray 1876 
Species
 axillaris
Pursh 1814
Species
 axillaris
Pursh 1814
Species
 microcephala
Nuttall 1841
Species
 angustifolia
Nuttall 1836
Species
 asperifolia
Lessing 1830
Species
 asperifolia
Lessing 1830
Species
 microcephala
Nuttall 1840
Species
 angustifolia
Nuttall ex De Candolle 1836
Species
 asperifolia
Lessing 1830
Species
 texensis
Jackson 1960
Subspecies
 angustifolia
(Nutt. ex DC.) Turner 2009
Species
 asperifolia
Lessing 1830
Species
 corbinii
Turner 2009
Species
 corbinii
Turner 2009
Subspecies
 asperifolia
(Lessing 1830) Turner 2009
Subspecies
 latior (Shinners)
(Jackson 1960
 I. texensis
) Turner 2009
Subspecies
 angustifolia
(Nutt. ex DC.) Turner 2009
Species
 asperifolia
Lessing 1830
Subspecies
 asperifolia
Lessing 1830
Subspecies
 latior (Shinners)
(Jackson 1960
 I. texensis
) Turner 2009
Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Species
 cheiranthifolia
H.B.K. 1820
Genus
 Iva
Linnaeus 1753R. C. JacksonJackson 1960Species
 annuaLinnaeus 1753Subspecies
 caudata
(Small 1899) Jackson 1960
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Subspecies
 caudata
(Small 1899) Jackson 1960
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Species
 frutescens
Linnaeus 1753Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria
(Bartlett) Jackson 1960
   Species
 hayesiana
Gray 1876 
Species
 frutescens
Linnaeus 1753Subspecies
 annuaJackson 1960
Genus
 Cyclachaena
Fresenius 1838
Species
 xanthifolia
(Nuttall 1818) Fresenius 1838 
Species
 xanthifolia
Nuttall 1818
Genus
 Chorisiva
(Gray 1884) Rydberg 1922
Genus
 Chorisiva
(Gray 1886) Rydberg 1922
Species 
nevadensis
(M. E. Jones 1883) Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Species 
dealbata(Gray 1852) Rydberg 1922
Species 
dealbataGray 1852
Species
 ambrosiaefolia
(Gray) Bentham & Hooker 1873
Species
 pedicellata
Rydberg 1922
Species
 lobataRydberg 1922
Sect.
 Linearbractea
Jackson 1960
Species
 microcephala
Nuttall 1840
Species
 angustifolia
Nuttall ex De Candolle 1836
Species
 asperifolia
Lessing 1830
Species
 texensis
Jackson 1960
Sect.
 Rhizoma
Miao et al.1995
Sect.
 Cyclachena
Gray 1884
aSect.
 Iva
Gray 1884
Genus
 Iva
Linnaeus 1753Miao, Turner & Mabry
Miao 
et al.
 1995a, b, &c
Species
 annuaLinnaeus 1753Subspecies
 annuaJackson 1960
dSpecies 
nevadensis
M. E. Jones 1883
Species
 acerosa
(Nutt.) Jackson 1960
Species
 ambrosiaefolia
(Gray 1852) Benth.&Hook. 1873
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 xanthifolia
(Nuttall 1818) Fresenius 1838
Species 
dealbata(Gray 1852) Rydberg 1922
Genus
 Cyclachena
Fresenius 1838
Species 
nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
Genus
 Oxytenia
Nuttall 1848
Genus
 Chorisiva
(Gray 1886) Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Species
 ambrosiaefolia
(Gray 1852) Benth.&Hook. 1873
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 xanthifolia
(Nuttall 1814) Fresenius 1838
Species 
dealbata(Gray 1852) Rydberg 1922
Genus
 Cyclachena
Fresenius 1838
Species 
nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
fiThe section 
Cyclachaena
 of Iva
 s.l. 
will be dealt within a later paper
because of its much closer relation-

ship to the other genera in the 

subtribe 
Ambrosiinae
.fl--Miao 
et al. 
1994. Analysis of section 

Cyclachaena
 of Iva
 s.l. was 
published in 1995. 
fiOur data suggest that 

I. ambrosiaefolia 
should be 

in a monotypic genus.fl
--Miao 
et al. 1995** Although Fernald and 
Griscom (1935) first established 

oraria 
as a subspecific 

designation of 
Iva frutescens
, 
it is a variety.  Jackson (1960)

specifically refers to his sub-

specific designation as a
subspecies.
   Carrington finds 
Iva annua 
caudata 
in synonymy with 

I.  a. 
annua. Paper in rewrite 
after review. See page 

XX, chapter 2.
B. L. Turner proposed (Turner 

2009a) that 
Iva corbinii
 was most 
closely related to 
I. axillaris, 
but with no colonial or rhizomatous 
features, and no evidence it is 

perennial, it's presence in 
section
 
Rhizoma
 should be reviewed.

--(PHC)Species
 axillaris
Pursh 1814
Species
 cheiranthifolia
H.B.K. 1820
Subspecies
 caudata
(Small 1899) Jackson 1960
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria
(Bartlett) Jackson 1960
  Species
 hayesiana
Gray 1876 
Species
 frutescens
Linneaus 1753Sect.
 Linearbractea
Jackson 1960
Species
 microcephala
Nuttall 1840
Sect.
 Rhizoma
Miao et al.1995
Sect.
 Iva
Gray 1884
Genus
 Iva
Linneaus 1753Miao, Turner & Mabry
Miao 
et al.
 1995a, b, &c
combined with 
Turner 2009a & 2009b
Species
 annuaLinneaus 1753Subspecies
 annuaJackson 1960
Species
 corbinii
Turner 2009
Subspecies
 angustifolia
(Nutt. ex DC.) Turner 2009
Species
 asperifolia
Lessing 1830
Subspecies
 asperifolia
Lessing 1830
Subspecies
 latior (Shinners)
Turner 2009  
[Jackson 1960
 I. texensis
]Genus
 Oxytenia
Nuttall 1848
Genus
 Chorisiva
(Gray 1886) Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Species
 ambrosiaefolia
(Gray 1852) Benth.&Hook. 1873
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 xanthifolia
(Nuttall 1814) Fresenius 1838
Species 
dealbata(Gray 1852) Rydberg 1922
Genus
 Cyclachena
Fresenius 1838
Species 
nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
Species
 axillaris
Pursh 1814
Species
 cheiranthifolia
H.B.K. 1820
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria
(Bartlett) Jackson 1960
   Species
 hayesiana
Gray 1876 
Species
 frutescens
Linneaus 1753Sect.
 Linearbractea
Jackson 1960
Species
 microcephala
Nuttall 1840
Sect.
 Rhizoma
Miao et al.1995
Sect.
 Iva
Gray 1884
Genus
 Iva
Linneaus 1753Species
 annuaLinneaus 1753Subspecies
 annuaJackson 1960
* Curiously, Asa Gray
 (1884) makes 
   a typographic change in the 

   name Iva xanthifolia 
as    I. xanthiifolia. 
This seems to have 

   been randomly disregarded by 
   subsequent authors, and 

   discarded
 in Jackson 1960.   Š(PHC)Asa Gray
 1884, placed  
Iva monophylla 
Walter 
into
Ambrosia 
as a synonym of

A. artemisiaefolia.

Š(PHC)Rydberg
 1922, lists

Ambrosia monophylla 
(Walt.) Rydberg 
as a new combination in 
Ambrosia, 

separate from 
A. artemisiaefolia 

(a name that is henceforth often 

spelled: 
A. artemisiifolia
). However, 

from here forward, 
Ambrosia 
monophylla is included within A. artemisiifolia.

Š(PHC)Iva monophylla
 Walter 1788

Was synonymized with

Ambrosia paniculata 
in 
De Candolle, 1836
.
   Š(PHC)cThe Taxonomic History of the Genus 
Iva
 (Asteraceae)
c  Cyclachena pedicellata 
Rydb. is
       given varietal status as 
I. xanthifolia

     var. 
pedicellata 
(Rydb.) Kittell 1941.

     Jackson (1960) synonymizes

     Cyclachena xanthifolia 
and
     C. pedicellata 
and I. x. 
var. 
pedicellata

      as I. xanthifolia 
within section

     Cyclachena 
of genus Iva.
d  Jackson places genera
      Leuciva 
Rydberg, 
Chorisiva

       Rydberg and 
Oxytenia

      Nuttall
 within section

      Cyclachena 
of genus Iva.

       All these genera restored

      by Miao, Turner, and 

      Mabry 1995.
Author
Genus
Subspecies
#Notes
Section
Species
b  Humboldt, Bonpland
      and Knuth are usually
      referred to as H.B.K. in

      contemporary citations.
f  Nuttall
 (1841) describes 
Iva foliolosa
 
largely 
on the basis of involucral 
bracts, and invoking the 
synonymy of 
Iva axillaris 
 Hooker, 1840 but it is a 
synonym 
of Iva axillaris 
in Asa Gray 
(1884).Iva paniculata
Nuttall 1841
Nuttall
 (1840) describes 
Iva paniculata
 largely 
on the basisof separate ‚male and female™
capitulae, it is a synonym 
of 
Iva xanthiifolia 
in Asa Gray (1884
).Asa Gray 
(1852) places species 

Iva xanthifolia
, in 
Euphrosyne
 and  
describes
 Euphrosyne 

ambrosiaefolia. 
Bentham and Hooker (1873) place

Euphrosyne
 xanthifolia
 and  
E. ambrosiaefolia 
in Cyclachaena.

Gray (1884) makes 
Cyclachaena 
asection of 
Iva
 instead of a genus.
E. L. Greene
 1902, designates a
type spec. coll. by C.  F.  Baker as

Iva obovata, 
but never publishes. 

It is synonymized by Jackson 1960

with I. axillaris. 
Iva cheiranthifolia 
 H.B.K. 1820, 

and I. asperifolia 
Lessing 1830 
were not included in Gray 1884. 
Perhaps it was because they were 

not known outside the islands

of the Caribbean and Mexico, 

and therefore not considered 

North American at the time. 
Kuntze
 1891, describes three 

subspecies of 
I. axillaris, 
I. a. normalis
, I. a. brevifolia, 
and I. a. linearifolia. 
They are all 

synonymized by Rydberg 1922

with I. axillaris. 
Carrington 2015 
Dissertation (not yet
in publication) including
all known contrib. to 2015.
Species
 corbinii
Turner 2009
IIIII-AIIIIVVVIVI-AVIIVIII   Doubtful Species:

Iva connata 
Sessé & Mocina
 
1887-1893. From Mexico.

R. C. Jackson found no 

evidence of type specimens.
Iva acuminata 
Nuttall
This is the title of a herbarium

sheet (PH00016157) in JSTOR 

Plants, but no such species was 

ever described. There is some

chance that a label with the 

words ‚acuminate -Nutt™ was 

misunderstood.  -PHC
Wildenow redescribes 
Iva annua

as Iva ciliata 
based upon 
Linneaus's mis-assignment 

of Iva annua 
to South America, 

believing it therefore to be a 
new northern species.
bbeea  Jackson restores the name 
      Iva annua 
based upon Linneaus'

     error and no specimens known 

     from South America.
Genus
 Oxytenia
Nuttall 1848
Species
 acerosa
Nuttall 1848
Figure 2-1 The Taxonomic History of the Genus 
Iva
The bold Roman numerals at the top of the main columns indicate the 
works of key authors in assembling the inventory of 
Iva 
at the times of their 
publications. The light blue ˚elds underneath each Roman numeral include the 
taxa that appear in the particular publication cited in the navy blue box at the 
top. The boxes that include the names of various taxa are color-coded to repre
-sent: genera, maroon; sections, orange; species, yellow; and subspecies, green.
#NOTE:  the category of  ‚box™ labelled subspecies includes all the subspeci˚c 
taxonomic categories used in the history of genus 
Iva. 
This includes subspecies, 
variety, and the use of lower case Greek letters to signify a variety. The term 
forma, does not appear to have been employed in 
Iva 
taxonomic history.
33(Figure 2-1, cont™d) Genus
 Iva
Linnaeus 1753Carl Linneaus
Linnaeus 1753Species
 annuaLinnaeus 1753Species
 frutescens
Linnaeus 1753Genus
 Iva
Linnaeus 1753Genus
 Euphrosyne
De Candolle 1836
Augustin P. de Candolle
De Candolle 1836
Species
 annuaLinnaeus 1753Species
 frutescens
Linnaeus 1753Genus
 Iva
Linnaeus 1753Genus
 Oxytenia
Nuttall 1848
Sect.
 Iva
Gray 1884
Sect.
 Iva
Gray 1884
Sect.
 Linearbractea
Jackson 1960
Asa Gray
Gray 1884
Species
 ciliataWilldenow 1804
Species
 ciliataWilldenow 1804
Species
 imbricata
Walter 1788
Species
 imbricata
Walter 1788
Species
 monophyllaWalter 1788
Species
 frutescens
Linnaeus 1753Species
 Hayesiana
Gray  1876
Sect.
 Cyclachaena
(Fresenius) Gray 1884
Species
 xanthiifolia*
Nuttall 1818
Species
 xanthifolia*
Nuttall 1818
Sect.
 Chorisiva
Gray 1884
Species 
Nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
Genus
 Oxytenia
Nuttall 1848
Genus
 Oxytenia
Nuttall 1848
Species
 acerosa
Nuttall 1848
Species
 dealbataGray 1852
Species
 ambrosiaefolia
(Gray) Gray 1884
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 ambrosiaefolia
(Gray) Gray 1884
 Genus
 Cyclachaena
Fresenius 1838
Species
 xanthifolia*
Nuttall 1818
Iva nevadensis
M. E. Jones 1883
Iva foliolosa
Nuttall 1841
Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Iva dealbata
Gray 1852
Iva H
ayesiana
Gray 1876 
Subspecies
 pubescens
Gray 1874
Subspecies
 axillaris
(Pursh 1814) Gray 1884
Subspecies
 latifolia
De Candolle 1836
Species
 axillaris
Pursh 1814
Species
 axillaris
Pursh 1814
Species
 axillaris
Pursh 1814
Species
 microcephala
Nuttall 1841
Species
 microcephala
Nuttall 1841
Species
 angustifolia
Nuttall 1836
Species
 angustifolia
Nuttall 1836
Genus
 Iva
Linnaeus 1753Per Axel Rydberg
Rydberg 1922
Species
 ciliataWilldenow 1804
Species
 imbricata
Walter 1788
Species
 caudata
Small 1899
Species
 frutescens
Linnaeus 1753Species
 Hayesiana
Gray  1876
Species
 oraria
Bartlett 1906 
Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria **
(Bartlett) Jackson 1960
  Species
 axillaris
Pursh 1814
Species
 hayesiana
Gray 1876 
Species
 axillaris
Pursh 1814
Species
 axillaris
Pursh 1814
Species
 microcephala
Nuttall 1841
Species
 angustifolia
Nuttall 1836
Species
 asperifolia
Lessing 1830
Species
 asperifolia
Lessing 1830
Species
 microcephala
Nuttall 1840
Species
 angustifolia
Nuttall ex De Candolle 1836
Species
 asperifolia
Lessing 1830
Species
 texensis
Jackson 1960
Subspecies
 angustifolia
(Nutt. ex DC.) Turner 2009
Species
 asperifolia
Lessing 1830
Species
 corbinii
Turner 2009
Species
 corbinii
Turner 2009
Subspecies
 asperifolia
(Lessing 1830) Turner 2009
Subspecies
 latior (Shinners)
(Jackson 1960
 I. texensis
) Turner 2009
Subspecies
 angustifolia
(Nutt. ex DC.) Turner 2009
Species
 asperifolia
Lessing 1830
Subspecies
 asperifolia
Lessing 1830
Subspecies
 latior (Shinners)
(Jackson 1960
 I. texensis
) Turner 2009
Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Species
 cheiranthifolia
Humboldt, Bonpland, Knuth
 1820Species
 cheiranthifolia
H.B.K. 1820
Genus
 Iva
Linnaeus 1753R. C. JacksonJackson 1960Species
 annuaLinnaeus 1753Subspecies
 caudata
(Small 1899) Jackson 1960
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Subspecies
 caudata
(Small 1899) Jackson 1960
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Species
 frutescens
Linnaeus 1753Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria
(Bartlett) Jackson 1960
   Species
 hayesiana
Gray 1876 
Species
 frutescens
Linnaeus 1753Subspecies
 annuaJackson 1960
Genus
 Cyclachaena
Fresenius 1838
Species
 xanthifolia
(Nuttall 1818) Fresenius 1838 
Species
 xanthifolia
Nuttall 1818
Genus
 Chorisiva
(Gray 1884) Rydberg 1922
Genus
 Chorisiva
(Gray 1886) Rydberg 1922
Species 
nevadensis
(M. E. Jones 1883) Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Species 
dealbata(Gray 1852) Rydberg 1922
Species 
dealbataGray 1852
Species
 ambrosiaefolia
(Gray) Bentham & Hooker 1873
Species
 pedicellata
Rydberg 1922
Species
 lobataRydberg 1922
Sect.
 Linearbractea
Jackson 1960
Species
 microcephala
Nuttall 1840
Species
 angustifolia
Nuttall ex De Candolle 1836
Species
 asperifolia
Lessing 1830
Species
 texensis
Jackson 1960
Sect.
 Rhizoma
Miao et al.1995
Sect.
 Cyclachena
Gray 1884
aSect.
 Iva
Gray 1884
Genus
 Iva
Linnaeus 1753Miao, Turner & Mabry
Miao 
et al.
 1995a, b, &c
Species
 annuaLinnaeus 1753Subspecies
 annuaJackson 1960
dSpecies 
nevadensis
M. E. Jones 1883
Species
 acerosa
(Nutt.) Jackson 1960
Species
 ambrosiaefolia
(Gray 1852) Benth.&Hook. 1873
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 xanthifolia
(Nuttall 1818) Fresenius 1838
Species 
dealbata(Gray 1852) Rydberg 1922
Genus
 Cyclachena
Fresenius 1838
Species 
nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
Genus
 Oxytenia
Nuttall 1848
Genus
 Chorisiva
(Gray 1886) Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Species
 ambrosiaefolia
(Gray 1852) Benth.&Hook. 1873
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 xanthifolia
(Nuttall 1814) Fresenius 1838
Species 
dealbata(Gray 1852) Rydberg 1922
Genus
 Cyclachena
Fresenius 1838
Species 
nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
fiThe section 
Cyclachaena
 of Iva
 s.l. 
will be dealt within a later paper
because of its much closer relation-

ship to the other genera in the 

subtribe 
Ambrosiinae
.fl--Miao 
et al. 
1994. Analysis of section 

Cyclachaena
 of Iva
 s.l. was 
published in 1995. 
fiOur data suggest that 

I. ambrosiaefolia 
should be 

in a monotypic genus.fl

--Miao 
et al. 1995** Although Fernald and 
Griscom (1935) first established 
oraria 
as a subspecific 
designation of 
Iva frutescens
, 
it is a variety.  Jackson (1960)

specifically refers to his sub-
specific designation as a
subspecies.
   Carrington finds 
Iva annua 

caudata 
in synonymy with 
I.  a. 
annua. Paper in rewrite 

after review. See page 
XX, chapter 2.
B. L. Turner proposed (Turner 
2009a) that 
Iva corbinii
 was most 

closely related to 
I. axillaris, 
but with no colonial or rhizomatous 
features, and no evidence it is 

perennial, it's presence in 
section
 
Rhizoma
 should be reviewed.

--(PHC)Species
 axillaris
Pursh 1814
Species
 cheiranthifolia
H.B.K. 1820
Subspecies
 caudata
(Small 1899) Jackson 1960
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria
(Bartlett) Jackson 1960
  Species
 hayesiana
Gray 1876 
Species
 frutescens
Linneaus 1753Sect.
 Linearbractea
Jackson 1960
Species
 microcephala
Nuttall 1840
Sect.
 Rhizoma
Miao et al.1995
Sect.
 Iva
Gray 1884
Genus
 Iva
Linneaus 1753Miao, Turner & Mabry
Miao 
et al.
 1995a, b, &c
combined with 
Turner 2009a & 2009b
Species
 annuaLinneaus 1753Subspecies
 annuaJackson 1960
Species
 corbinii
Turner 2009
Subspecies
 angustifolia
(Nutt. ex DC.) Turner 2009
Species
 asperifolia
Lessing 1830
Subspecies
 asperifolia
Lessing 1830
Subspecies
 latior (Shinners)
Turner 2009  
[Jackson 1960
 I. texensis
]Genus
 Oxytenia
Nuttall 1848
Genus
 Chorisiva
(Gray 1886) Rydberg 1922
Genus
 Leuciva
Rydberg 1922
Species
 ambrosiaefolia
(Gray 1852) Benth.&Hook. 1873
Subspecies
 ambrosiaefolia
Jackson 1960
Subspecies
 lobata(Rydberg 1922) Jackson 1960
Species
 xanthifolia
(Nuttall 1814) Fresenius 1838
Species 
dealbata(Gray 1852) Rydberg 1922
Genus
 Cyclachena
Fresenius 1838
Species 
nevadensis
M. E. Jones 1883
Species
 acerosa
Nuttall 1848
Species
 axillaris
Pursh 1814
Species
 cheiranthifolia
H.B.K. 1820
Subspecies
 macrocarpa
(Blake 1939) Jackson 1960
Species
 imbricata
Walter 1788
Subspecies
 frutescens
Jackson 1960
Subspecies
 oraria
(Bartlett) Jackson 1960
   Species
 hayesiana
Gray 1876 
Species
 frutescens
Linneaus 1753Sect.
 Linearbractea
Jackson 1960
Species
 microcephala
Nuttall 1840
Sect.
 Rhizoma
Miao et al.1995
Sect.
 Iva
Gray 1884
Genus
 Iva
Linneaus 1753Species
 annuaLinneaus 1753Subspecies
 annuaJackson 1960
* Curiously, Asa Gray
 (1884) makes 
   a typographic change in the 
   name Iva xanthifolia 
as 
   I. xanthiifolia. 
This seems to have 

   been randomly disregarded by 
   subsequent authors, and 

   discarded
 in Jackson 1960.   Š(PHC)Asa Gray
 1884, placed  
Iva monophylla 
Walter 
into

Ambrosia 
as a synonym of
A. artemisiaefolia.
Š(PHC)Rydberg
 1922, lists
Ambrosia monophylla 

(Walt.) Rydberg 
as a new 
combination in 
Ambrosia, 
separate from 
A. artemisiaefolia 
(a name that is henceforth often 

spelled: 
A. artemisiifolia
). However, 

from here forward, 
Ambrosia 

monophylla is included within 
A. artemisiifolia.

Š(PHC)Iva monophylla
 Walter 1788

Was synonymized with

Ambrosia paniculata 
in De Candolle, 1836
.   Š(PHC)cThe Taxonomic History of the Genus 
Iva
 (Asteraceae)
c  Cyclachena pedicellata 
Rydb. is
       given varietal status as 
I. xanthifolia
     var. 
pedicellata 
(Rydb.) Kittell 1941.

     Jackson (1960) synonymizes

     Cyclachena xanthifolia 
and
     C. pedicellata 
and I. x. 
var. 
pedicellata

      as I. xanthifolia 
within section

     Cyclachena 
of genus Iva.
d  Jackson places genera
      Leuciva 
Rydberg, 
Chorisiva

       Rydberg and 
Oxytenia

      Nuttall
 within section

      Cyclachena 
of genus Iva.

       All these genera restored
      by Miao, Turner, and 
      Mabry 1995.
Author
Genus
Subspecies
#Notes
Section
Species
b  Humboldt, Bonpland
      and Knuth are usually

      referred to as H.B.K. in

      contemporary citations.
f  Nuttall
 (1841) describes 
Iva foliolosa
 largely 
on the basis of involucral 
bracts, and invoking the 
synonymy of 
Iva axillaris 

 Hooker, 1840 but it is a 
synonym 
of Iva axillaris 
in Asa Gray 
(1884).Iva paniculata
Nuttall 1841
Nuttall
 (1840) describes 
Iva paniculata
 largely 
on the basisof separate ‚male and female™
capitulae, it is a synonym 
of Iva xanthiifolia 
in Asa Gray (1884
).Asa Gray 
(1852) places species 
Iva xanthifolia
, in 
Euphrosyne
 and  
describes
 Euphrosyne 

ambrosiaefolia. 
Bentham and Hooker (1873) place
Euphrosyne
 xanthifolia
 and  E. ambrosiaefolia 
in Cyclachaena.

Gray (1884) makes 
Cyclachaena 
a
section of 
Iva
 instead of a genus.
E. L. Greene
 1902, designates a

type spec. coll. by C.  F.  Baker as
Iva obovata, 
but never publishes. 
It is synonymized by Jackson 1960
with I. axillaris. 
Iva cheiranthifolia 
 H.B.K. 1820, 
and I. asperifolia 
Lessing 1830 

were not included in Gray 1884. 

Perhaps it was because they were 

not known outside the islands

of the Caribbean and Mexico, 
and therefore not considered 
North American at the time. 
Kuntze
 1891, describes three 
subspecies of 
I. axillaris, 

I. a. normalis
, I. a. brevifolia, 
and 
I. a. linearifolia. 
They are all 

synonymized by Rydberg 1922

with I. axillaris. 
Carrington 2015 
Dissertation (not yet
in publication) including
all known contrib. to 2015.
Species
 corbinii
Turner 2009
IIIII-AIIIIVVVIVI-AVIIVIII   Doubtful Species:
Iva connata 
Sessé & Mocina
 
1887-1893. From Mexico.

R. C. Jackson found no 

evidence of type specimens.
Iva acuminata 
Nuttall
This is the title of a herbarium

sheet (PH00016157) in JSTOR 

Plants, but no such species was 

ever described. There is some

chance that a label with the 

words ‚acuminate -Nutt™ was 

misunderstood.  -PHC
Wildenow redescribes 
Iva annua

as Iva ciliata 
based upon 
Linneaus's mis-assignment 
of Iva annua 
to South America, 
believing it therefore to be a 
new northern species.
bbeea  Jackson restores the name 
      Iva annua 
based upon Linneaus'

     error and no specimens known 

     from South America.
Genus
 Oxytenia
Nuttall 1848
Species
 acerosa
Nuttall 1848
34is followed by fi(Nutt. in litt. 1825).fl To what De Candolle is referring in litt. 1825 is a mystery. All 
citations for I. angustifolia refer to the De Candolle Prodromus of 1836, not an 1825 publication.1838  J. B. Fresenius publishes the genus Cyclachaena (Fresenius 1838). On its surface, this ap-pears to be a very unusual publication. First, the name Fresenius appears nowhere in it or near it in Schlechtendal™s (series editor) massive collection. Second, there are no species names associated 
with this genus description. The collection is attributed to ‚principe Maximiliano, Neovidense™ that 
would be Prince Alexander Philipp Maximilian zu Wied-Neuwied, who collected along the up
-per Missouri river after 1832 through before 1840. It can be inferred that xanthifolia is the species involved because it is the only related species with no corolla (or reduced to a thin disc) associated 
this reference (e.g. Torrey and Gray 1842, Gray 1884, Rydberg 1922, Jackson 1960), but there are 

Iva (Jackson 1960), he appears to have cited the reference as appearing in 1836, as do many others; upon examination, 
1836 is in doubt with Wied™s expedition noted as from 1832-1834.
1840  William Jackson Hooker published 
I. axillaris 
in Flora Borealis-Americana 
(Hooker 1840), Volume I. When Nuttall (Nuttall 1840) describes 
I. foliolosa. Hooker™s variety of 
I. axillaris 
 is mentioned as a synonym. Asa Gray later places 
I. foliolosa as a synonym of I. axillaris (Gray 1884).1841  Thomas Nuttall™s 1841 publication, 
35

 (Nuttall 1841), included new species descriptions introducing Iva foliolosa, and I. paniculata. Nuttall references I. foliolosa as the ‚subspecies™ 
Iva axillaris,, Hooker. Although he 
distinguishes I. foliolosa from Pursh™s 
I. axillaris by leaf and involucre, it is placed as a synonym of I. axillaris in Asa Gray™s Synoptical Flora of North America, 1884.
Nuttall™s description of 
Iva paniculata includes the mention of separate ‚male and female™ capitula, 
Iva section with the note that I. xan-thifolia is ‚nearly allied to the preceding (I. paniculata). I. paniculata is placed as a synonym of I. xanthifolia (now spelled ‚xanthiifolia) in Asa Gray™s
 Synoptical Flora of North America, 1884.
Nuttall™s remaining new species in this publication is 
I. microcephala, 

-1848   Nuttall™s 

of Upper California (Nuttall 1848) contains the founding description of the genus and the 
-form in description. He also notes how similar this new genus is to 
and Cyclachaena. Lastly commenting that Cyclachaena should be reconsidered as an is placed as a section within Iva in Jackson™s revision (Jackson 1960, page 828) but restored as a genus 
in the genetically based work of Miao, Turner, and Mabry (Miao, Turner, and Mabry 1995).
1852  Asa Gray™s description, in 

 (Gray 1852) of is the origi-nal description of a species that still exists in today™s taxonomy although it is currently assigned to 
36the genus Leuciva, after being moved around as a species in Iva section Cyclachaena (Gray 1884, Jackson 1960) and after that, the sole species in the genus Leuciva (Rydberg 1922, Miao, Turner, 
and Mabry 1995). This work also sees Gray™s description of 

In Gray™s 
own 1884 Synoptical Flora of North America
, he places this species into genus Iva. It will end up in genus Cyclachaena, (Miao, Turner, and Mabry 1995) but its future there is uncertain.
1873  G. Bentham and J. D. Hooker in their version of , take Gray™s 
Euphrosyne 

and place it into genus Cyclachaena  (
) where xanthifo-lia is returned to as well (Bentham and Hooker 1873).1874  
1842, (Dupree 1988, pages 59-65 and 67-68), he turned down the expedition in July of the year it de-
-
to resign in April 1840. He was appointed Fisher Professor of Natural History at Harvard in 1842). 
With the return of the survivors of the expedition, and the remaining undamaged parts of the collec
-tions safely stored, Gray eventually published part of the botanical discoveries (Gray 1874) though parts 
United States Exploring 

Botany, in 1874. In it, he describes the variety Iva axillaris var. 
on the basis of its lax spread-ing hairs, from the Bay of San Francisco. It appears in his editions of the Synoptical Flora of North 
America, starting in 1884, but is synonymized with I. axillaris according to Rydberg (1922).
1876  Asa Grays™s description in the 

37(Gray 1876) of Iva Hayesiana (now I. hayesiana) from California, has the curious note, fiIn memory of the estimable discoverer, the late Mr. Sutton Hayes, whose specimens, however, were indetermin
-able, the heads having all fallen from their short peduncles.fl1883  Marcus E. Jones™s description of 
Iva nevadensis in the American Naturalist of a species he 

Asa Gray (1884) Places Iva Nevadensis (note new capitalization) in Iva section Chorisiva, for the -rolla.1884 
Synoptical Flora of North America (Column III in Figure 2-1) of which the second edition was published in 1886 (Gray 1884). Here he presents three divisions of genus Iva; namely Iva, Cyclachaena, and Chorisiva. The genus 
Nuttall is retained. The most critical diagnostic characters of the three sections are: Iva, evi
-Cyclachaena, very short or rudimentary Chorisiva
I. axillaris, Gray™s 1874 variety 
I. I. a. axillaris. Iva chei-ranthifolia  H.B.K. 1820, and I. asperifolia Lessing 1830 were not included in Gray 1884. Perhaps it was because they were not known outside the islands of the Caribbean and Mexico, and therefore not considered North American at the time.
1899 John K. Small published Iva caudata in 
States (Small 1899). He distinguishes this species from I. ciliata (now I. annua) by the fismoother 38Iva by Jackson (1960, page 812) he places I. caudata as a variety of I. annua; I. annua var. 
caudata. The data to formally synonymize I. a. caudata with I. a. annua  are presented by me on page (54).1902  E. L. Greene designates a specimen collected by C. F. Baker as the type specimen of 
Iva in a list of western plants (Greene 1902), but it is never published. It is formally made a synonym of I. axillaris by Jackson (1960, page 823). It does appear as a herbarium sheet in JSTOR 
Plants (the seal on this sheet reads, Herbarium of Pomona College 57543).1906  Harley H. Bartlett publishes his description of species Iva oraria (Bartlett 1906) based upon the magnitude of difference between the sizes of leaf, involucre, and achene from plants originating 
at the northern and southern ends of the U.S. Atlantic coast. Oddly he notes that intermediates occur 
at the Texas locations at the west end of the U. S. Gulf of Mexico coast. In 1935, it is placed as a 
variety of  Iva frutescens (Fernald and Griscom 1935). The revision of 
Iva by Jackson (1960) revises the status to subspecies as I. frutescens subsp. oraria (Jackson  1960, page 818).1922  This is the year of the publication of the New York Botanical Garden™s 
North American Flora, Volume 33, part I
 (Column IV in Figure 2-1) that includes the Ambrosiaceae by Per Axel 
Rydberg. In the announcement on the frontispiece, North America is taken to include fiGreenland, 
Central America, the Republic of Panama, and the West Indies, except Trinidad, Tobago, and Cura
-
This work presents I. cheiranthifolia H.B.K. and I. asperifolia Lessing in the context of their status in genus Iva, 
39Rydberg does away with the three sections of 
Iva (sections Iva, Cyclachaena, and Chorisiva) that were established by Gray (1884) by reestablishing the generic status of Cyclachaena, as well as el-evating the section Chorisiva by Gray (1884) into a genus in its own right. This makes all the section 
levels of Iva disappear because section Iva is not valid standing alone. In addition, he describes the new genus Leuciva Rydberg 1922, as the monotypic genus ‚home™ of the former 
Iva sec. Cyclachae-Gray; yielding (Gray 1852) Rydberg 1922, based largely on the lack of 
  retains this status to this day.
In the intervening 36 years since Gray™s 1886 
Synoptical Flora of North America
, second edition, both Iva caudata Small and Iva oraria Bartlett were described and they both appear in genus Iva in Rydberg 1922. In addition, Rydberg himself makes two additional descriptions to the genus 
Cy-clachaena; C. pedicellata, and 1939  
cultivation of (then called) Iva ciliata by the American Indians of the Middle Archaic to the Late 
Mississippian periods, designates the large-seeded cypselae recovered as part of archaeology as 
belonging to a new variety, 
I. ciliata var. 
macrocarpa. 
In this paper he also states his opinion (Blake 1939, page 85) that the extremities of the length to width ratio Small (1899) used as the principle criterion to distinguish (his then species) Iva caudata from I. annua will turn out to be points on a 1941  Sister Teresita Kittell of Holy Family College, Manitowoc, Wisconsin, in 
A Flora of Arizona 
and New Mexico, that she co-authored with Ivar Tidestrom (Tidestrom and Kittell 1941), placed 
Cy-40clachaena pedicellata 
C. xanthifolia; namely C. xanthifolia pedicellata 
1960  This year marks the publication of A Revision of the Genus 
Iva L. by R. C. Jackson (Column V, Figure 2-1). As a graduate student, R. C. Jackson had worked under Charles B. Heiser. In a 2005 
conversation with me (pers. com.), Heiser remarked that he had been interested in Iva and its role in history and that he had tried over the years to interest various graduate students to pursue genus Iva further, but that Jackson was the only one who made an effort in that direction (Jackson 1960). 
One of the most important changes in Iva taxonomy in this work was the recognition of the syn-onymy of Iva ciliata Willd., with 
Iva annua L. This meant that the variety 
I. ciliata var. 
macrocarpa, 
became a new combination; I. annua var. 
macrocarpa. 
Jackson placed Small™s (Small 1899) species 
I. caudata at varietal rank under I. annua as I. a. var. 
caudata. He also placed Bartlett™s species 
I. oraria, that Bartlett tried at length to justify separating from I. frutescens, I. frutescens, as I. f. var. 
oraria. Jackson proposed an inclusive approach to genus Iva, wherein he took the species previously housed in the genera Cyclachaena, Leuciva, Chorisiva, and and conjoined them all into an expanded version of section Cyclachaena, within Iva. Rydberg™s 
Cyclachaena pedicellata that had Cyclachaena xanthifolia pedicellata (Rydberg) Kittell) by Kittell 
(Tidestrom and Kittell 1941) and the species 
Cyclachaena xanthifolia 
were all synonymized to be Iva xanthifolia Nuttall in section Cyclachaena. The species that had resided in genus 
Iva, sensu Rydberg 
Iva,  based on bract morphology, and 
Rhizoma, with the sole occupant I. axillaris, apparently based on the colonial rhizomatous habit.41Jackson does describe one new species in this revision, Iva texensis, from the same branch of his phylogenetic interpretation (Jackson 1960, page 843) as I. microcephala, I. angustifolia, I. asperifo
-lia, in the n-16 chromosome group. He seems to have an uncertainty about some aspects of this spe-cies. He designates a type and a co-type but then lists a number of specimens that he declares fiin in-volucral length and plant height they are not always in agreement with the type.fl He then refers to 7 coastal Texas specimens as fiSpecimens tentatively assigned to
 I. texensis.fl In hindsight, this almost seems like a premonition of B. L. Turner™s placing of 
I. texensis I. asperifolia, (Turner 2009a) where its name reverts to 
I. asperifolia var. 
latior Shinners (1964). Had Jackson™s 
species, Iva texensis been ultimately recognized as a species, it would have been (correctly) named I. texensis.1964  Lloyd H. Shinners in a rather heterogeneous paper New Names and Records for Texas Com
-positae, describes a new variety, 
Iva angustifolia (that is misspelled ‚augustifolia™) var. 
latior. This 
latior).1995  In 1995, B. Miao, B. L. Turner, and T. J. Mabry published three papers that brought the 
taxonomy of the subtribe Ambrosiinae into the age of molecular taxonomy (Miao, Turner and Mabry 
1995a, b, and c). In order they are: a)  Iva (Asteraceae, Heliantheae) 
, b) 
. Cyclachaena of 
 Ambro-siinae . Of these publications, the second (Miao, Turner and Mabry 1995b) 
has the most direct implications for the taxonomic structure of genus Iva and its close relatives (Fig-42ure 2-1, column VI).
Their analysis of cpDNA implies that the portion of Jackson™s genus 
Iva represented by sections Iva and  are basically monophyletic and sound. They are comprised of n=16 species, 
() and n=17 species (Iva). Jackson™s section 
Cyclachaena is paraphyletic and its divi-sion into separate genera, and separate from Iva synapomorphies that would unite the 5 species from Jackson™s section 
Cyclachaena. All the section 
Cyclachaena species form a category of n=18 chromosomes. In their analysis of the Wagner and 
Dollo trees that they produced to evaluate the way these relationships could be handled, they offer 
a couple scenarios that would satisfy most of the incongruities (Miao, Turner and Mabry 1995a, b). 
The one I felt they emphasized was to reestablish the ‚small genus™ system begun by Rydberg (1922). 
This would revive the genera and Euphrosyne. 
This genera (except for Cyclachaena that would have two species, 
and C. xanthifolia) that might obscure that many similarities between them, and alternatively, revive and expand the 
formal genus Cyclachaena to include the species formerly housed in the genera Euphrosyne
 and , without which, Cyclachaena would be a paraphyletic group. The phylogenies constructed 
from the cpDNA shows 
Cyclachaena is more closely related to Euphrosyne
 and than to the ‚proper™ n=16, and n=17 members of 
Iva s. str. These are the relationships I have depicted in Figure 2-1 Column VI, except that I have not included the genera 
Euphrosyne 
and 2006  John L. Strother™s entries in the Asteraceae within the Flora of North America, Volume 21 
43(Strother 2006), include Iva and related genera. In this part of the work, he ignores Iva asperifolia, but he does place in synonymy under I. angustifolia both I. texensis Jackson, and I. angustifolia var. 
latior Shinners.2009  In this year there were two changes to genus Iva sensu strictu., both in publications by Billie L. Turner, of the Plant Resources Center of the University of Texas at Austin. 

I. asperifolia, I. angustifolia, and I. angustifolia, var. 
latior Shinners (formerly I. texensis Jackson). By gaining access to impor-tant specimens of I. asperifolia from the major portion of its range in northeast Mexico, he is able to demonstrate a case for placing all three in synonymy. Because the senior name in the group is 
I. asperifolia, Lessing 1830, the other names became the junior synonyms resulting in the species I. as-perifolia being the correct species name, and its three varieties being; I. a. Lessing var.
 asperifolia, I. a. var. 
angustifolia (Nutt. Ex DeCandolle) Turner
, and I. a. var. 
latior (Shinners) Turner
. This brings 
the status of Iva names to the appearance of column VII, Figure 2-1.
In Billie Turner™s second paper (Turner 2009b) he describes the new species 
from the islands of the Colorado River bottoms of Travis county Texas (not the larger more famous Colorado 
River). It is described from a small population on a few islands and is singular for its leaf-like sub-Robert Corbin, the eponymous collector, picked a specimen for Turner to evaluate, was abducted 
from the location in the interim time, leaving just a hole.  Turner considers this species close to 
Iva axillaris, but it has taproots, no rhizomes and is reported to have n=16 chromosomes.2015  Iva caudata, Small, 44whose range is overlapped virtually completely with the range of Iva annua, that was later placed as a variety of  I. annua L. namely I. a. var. 
caudata (Small) Jackson, should be placed in synonymy with I. a. var. 
annua. The evidence for this comprises the bulk of chapter 4. This brings the discus
-
genus Iva. In the case of the I. asperifolia species complex discussed above, there are three appar-ent variations that seem to intergrade from one variety to the other where they contact each other 
across the extent of its natural range. In the case of the extant range of I. annua, the two extremes in appearance, bract length to width ratios, are nested within the same geographical range as well as the intergrades between them. On this basis, I would suggest that there is a variety comprised of the 
giant-seeded specimens from archaeological discovery, 
I. a. macrocarpa
, and the sole variety for the extant populations of I. annua, namely I. a. annua, and that I. a. caudata be considered a synonym of I. a. annua (See Chapter 3).It is fortunate that this genus has attracted enough interest to have so many of its issues illuminated 
e.g. Solidago, andhoused within the Asteraceae
 45REFERENCES46REFERENCESBARTLETT, H. H. (1906). fiThe Salt-Marsh Iva of New England.fl Rhodora 8(86): 25-26.BENTHAM, G., HOOKER, J. D. (1873). . London, Lovell, Reeve & Co.BLAKE, S. F. 
(1939). fiA New Variety of Iva Ciliata From Indian Rock Shelters in the South-Central 
United States.fl Rhodora 41(483): 81-86.CLA
USEN
, R. 
T. (1941). 
fiOn the use of the terms fiSubspeciesfl and fivarietyfl.fl Rhodora 43(509): 157-167.
DE CANDOLLE, A. P. 
(1836). 
. Paris.DUPREE, A. 
H. (1988). 
. Baltimore, Johns Hopkins University Press.FERNALD, M. L. G., LUDLOW (1935). fiThree Days of Botanizing in Southeastern Virginia.fl 
Rhodora 37(437): 167-189.FRESENIUS, J. B. (1838). fiSemina in Horto Bontanico Frankefurtensi 4.fl Linnaea 12: 78.GRAY, A. 
(1852). 



 Washington, Smithsonian Institution.
(1874). Botany. 


, C. Sherman. 17: 350. (1876). fiMiscellaneous Botanical Contributions.fl 
Arts and Sciences III: 78.(1884). Synoptical Flora of North America
. New York, Smithsonian Institution.
 (1886). Synoptical Flora of North America
. Washington, D. C., Smithsonian Institution.
GREENE, E. L. (1902). 
. Stanford, California, Stanford 
 University.
HAMILTO
N, C. W., R
EICHARD, SARAH H. (1992). fiCurrent Practice in the Use of Subspecies, Variety, 
 
Taxon 
41(3): 485-498.HOOKER, W. J. 
(1840). Flora Boreali-Americana
 London, Treuttel, Jun. & Richter.
47HUMBOLDT, A., B
ONPLAND, KUNTH (1820). 
. Paris.JACKSON, R. C. (1960). fiA revision of the genus 
Iva L.fl University of Kansas Science Bulletin 41: 793-876.JARVIS, C. (2007). 
 London, The Linnean 
Society of London.JONES, M. E. (1883). fiNew Plants from California and Nevada, etc. II.fl 
American Naturalist 17(9): 973.LESSING, C. F. 
(1830). fiIn D. F. L. von Schlechtendal and L. A. von Chamisso L.K.A von: Plantarum 
Mexicanarum.fl Linnaea 5: 151.LINNAEUS, C. (1753). . Holmiae [Stockholm], Laurenti Salvii.
MATASYOH, J. C., KIPLIMO, JOYCE J., KRUBIU, NICHOLAS M., HAILSTO
RKS, TIFFAN
Y P. 
(2007). fiChemical composition and antimicrobial activity of essential oil of Tarchonanthus camphoratus.fl 
Food Chemistry 101: 1183-1187.
MIAO, B., TURNER, B., MABRY, T. (1995a). fiMolecular phylogeny of 
Iva (Asteraceae, Heliantheae) based on chloroplast DNA restriction site variation.fl 
 195: 1-12.  (1995b). fiChloroplast DNA variations in Sect. 
Cyclachaena of Iva (Asteraceae).fl American  82(7): 919-923.  (1995c). fiSystematic implications of chloroplast DNA variation in the subtribe 

(Asteraceae: Heliantheae).fl  82(7): 924-932.NUTTALL, T. (1818). 
. Philadelphia, D. Heartt. (1841). fiDescriptions of new Species and Genera of Plants in the natural Order of the COM-
and a Visit to the Sandwich Islands and Upper California, during the Years 1834 and 1835.fl 

 7: 283-453. (1848). fiDescriptions of Plants Collected by Mr. William Gambel in the Rocky Mountains 
and Upper California.fl 
 III: 7-26.PURSH, F.
 (1814). 

. London, White, Cochrane, and Co.
RYDBERG, P. A. 
(1922). Tribe 1. Iveae. 
North American Flora
. New York, New York Botanical Gar
-den. 33: 3-13.48SHINNERS, L. H. (1964). fiNew Names and Records for Texas Compositae.fl 
Sida 1(6): 373-388.SMALL, J. K. (1899). fiUndescribed Species from the Southern United States.fl 

 1: 278-290.STROTHER, J. L. (2006). Iva. Flora of North America North of Mexico
. F. o. N. A. E. 
 Committee. New York, Oxford University Press. 
21: 25-28.TIDESTROM, I., KITTELL, SISTER TERESITA (1941). A Flora of Arizona and New Mexico
. Washington, D. 
  C., Catholic University of America Press.
TORREY, J., M.D., GRAY, A
SA, M.D. (1842). Flora of North America
. New York, Wiley & Putnam.
TOURNEFORT, J. P. 
(1700). 
. Paris, Parisiis Etypographia Regia.TURNER, B. L. (2009a). fiTaxonomy of 
Iva angustifolia and I. asperifolia (Asteraceae).fl  91(1): 76-83.  (2009b). fi(Asteraceae): A Remarkable New Species from Travis County, 
Texas.fl 
Lundellia 12: 5-7.VAILLANT, S. (1719). ރtablissement

 (1719): 277-318.WALTER, T. (1788). IVA. 
Flora Caroliniana
. London, Sumptibus J. Fraser: 232.WILLDENOW, C. L. (1804). IVA. 
. Berlin. III, pt. 3: 2386-2387.49CHAPTER 3The Taxonomic History of 
Iva annua (ASTERACEAE)AbstractThe Asteraceae species, 
Iva annua, Iva by Linnaeus in his 
included erroneous location data that led to Iva annua soon being redescribed under the name Iva ciliata. This chapter reviews the taxonomic history of the species, before and after Linnaeus, the 
status of the varieties that have been published since 1753, and proposes a revision. Iva annua var. 
macrocarpa, 
is the variety erected to contain the large-fruited specimens that were discovered in ar
-than ever comprising a wild form of the modern species (Blake 1939). A fourth variety, 
Iva ciliata var. 
latifolia  (de Candoll 1836) was short lived, and appears to be identical to Iva annua var. 
annua. Introduction
Iva annua, is arguably the most well-known, important, and controversial species in genus 
Iva. Discovery of its cached seeds at archaeological sites in the central United States, in the early twenti-eth century, propelled it to notice as an agricultural entity in the early development of the food seeds, 
in what is known as the Eastern Agricultural Complex (Harrington 1924; Gilmore 1931; Smith 1992,  
Wagner and Carrington 2014). 
50The  of Linnaeus and the Linnaeus 1754, and 1753 respectively) constitute the starting point for genera and species respectively, for all modern 
plant taxonomy [Article 13, 
], including the genus Iva,  I. annua, and I. frutescens. As these volumes [and ] by Linnaeus are the beginning of the modern system of naming plant species, names published before that time have no standing in modern nomenclature unless published again later under the rules of nomenclature.Pre-Linnaean & Linnaean History
An early (pre-Linnaean) introduction to the species that would ultimately be described as Iva an-nua, was by Sébastien Vaillant, in his 


 (page 310), under the name Tarchonanthos 
(Vaillant 1719). 
Tarchonanthus 
(see account starting on page 16).Vaillant™s species diagnosis of the American 
Tarchonanthus 
is very short: Tarchonanthos folio tri
-[Tarchonanthus
 (modern spelling) with three veined, toothed leaves, 
T. camphoratus; 
the African 
species has the mid-vein as the only prominent leaf vein and a non toothed margin, and although 

plant are upright, not pendulous. Vaillant ends with what would be an early example in a series of er
-roneous location notes throughout the history of I. annua, in the note: . There are no 
known Iva species native to South America (Jackson 1960). I have found no evidence for how any 
51Iva specimens were obtained by these earliest writers. The implication is they were delivered from an 
unknown collector in the late 17th or early 18th century who visited southeast North America and Peru.
In 1737, Linnaeus published the Hortus Cliffortianus (Linnaeus 1737). Here he placed the discus-sion of Tarchonanthus 
under the name  [PARTHENIUM leaves lanceolate with teeth.] and 
Adriani van Royen, physician and botanist born in Leiden in 1704, prevailed on Linnaeus (Gorton 1847; Thijsse 2013) to spend some time with him in the preparation of 

-mus that appeared in 1740 (van Royen 1740). Here Tarchonanthus 
is directly cited from Vaillant 
1719; only the African species is described in the main text (page 152); the American species is 
relegated to the Stirpes Vagae, 
uncertain plants (page 538).In 1748 Linnaeus published the Hortus Upsaliensis.name Iva (Linnaeus 1748, page 285). It followed time, and cites the Stirpes Vagae
 page from van Royen [
] but from the details of the text (Radix annua noted), he seems to be describing Iva annua. Linnaeus never gave an explanation for the name Iva; it is widely believed to have been adapted from the species Ajuga iva (Lamiaceae) because of its similar odor (Austin 2004, Correll &Correll 1982, Hickman 1993, Diggs et al. 1999), fisome medicinal plantfl (Fernald 1950, Weber 1987).
When Iva species appeared by Linnaeus in 1753, the genus Iva had already been described in Hor-tus Upsaliensis (Linnaeus 1748). The text is translated below (translation from Latin and parentheti
-cal inclusions by me).52I. IVA
 leaves lance-ovate, serrate, stem annual. Tarchonanthus
 cordate leaves serrated with three main veins. B. Jussiaei 
  
Habitat in America.  
Reared in a greenhouse, blooming late, annual. Description. Root annual. Stem human height, erect, striated, hairs white sparsely scattered. Branches few. Lower leaves opposite, upper leaves alternate, 
lance-ovate, acuminate, petiolate, teeth somewhat large, three main veins, rough. 
leaves (bracts). Flowers nodding as in species 1 in Hortus Cliffortianus but female 
The species of marshelder known today as Iva annua 
species in the genus Iva described by Carl Linnaeus in (Linnaeus 1753). The bot
-tom of page 988, positioned just above the continuation on page 989 are shown next page (Figure 3-1).
Code(s) of Botanical Nomenclature, were not in place at the birth of modern binomial nomenclature. In Jackson™s revision of the genus 
Iva (Jackson 1960, page 808), he says he examined the type, from the Herbarium of the Linnean Society of London by photograph, and that his illustrations, Jackson 1960, 12-16, on page 845) were drawn from this ‚type™, (Figure 3-2). According to Jarvis in Order 
53out of Chaos Linnaean Plant Names and their Types, (Jarvis 2007) Jackson designated a specimen 
from the Herbarium of the Linnean Society of London (Herb. Linn. No. 1116.1 (LINN) as the lec
-totype in his revision (Jackson 1960, page 808). In the online resource JSTOR Plant Types, what is 
labeled as this specimen is called the lectotype and occupies a single sheet with virtually no date or Figure 3-1 
 Linnaeus™s original 
Iva 
species
 descriptions in Species Plantarum 
Species Plantarum, by . The Iva. Iva annuaIva frutescens. Scanned 
collection information (Figure 3-3). It is the only one of JSTOR™s several 
Iva annua digitized speci-men sheets designated as a lectotype, and in all important respects seems to be consistent with the illustrations in Jackson™s reorganization of 
Iva (Jackson, 1960, page 845).Linnaeus™s 1753 diagnosis for 
Iva annua, is composed of very few words (translation by me), fiIVA
 leaves are lanceolate-ovate, on an herbaceous stem. Tarchonanthus
 leaves bearing three main veins. 54Figure 3-2 
 Jackson™s illustration (1960) of Linnaeus possible lectotype of 
Iva annua 
These 
A Revision of the Genus 
Iva annua 



55Figure 3-3 
 Lectotype of 
Iva annua 
L. from the Linnean Society of London
 The specimen sheet 
Iva annua L. from the Linnean 


that were drawn from this specimen.
56Roy. lugbd, 538. Habitat in South America. D. B. Jussieu,fl followed by the symbol for the sun, an 
indication that the species is an annual (Stearn 1962).Linnaeus™s mis-characterization of 
Iva annua as a South American species would start over two 
centuries of confusion and result in a long-unresolved synonymy. Throughout 
, Linnaeus often added notes about range or collection data. Unfortunately, the entry for 
Iva annua reads, fi Habitat in America meridionalifl. The Latin term ‚meridionali™ is generally taken to mean 
south or southern, as in South or Southern America. This Latin word, reportedly dating from the 
fourteenth century, has occasionally been interpreted as meaning ‚on or along a meridian.™ The case 
for interpreting ‚meridionali™ as south is consistent with his other comparable term, ‚australis™ that he 
seems to intend to be read as southern, as when he writes, fi
fl Linnean scholar W. T. Stearn writes in his 


(Stearn 1957) that Habitat in America meridionali, especially when considering collections made by Plumier, referred to locations made in the West Indian Islands, 
I. annua, credits D. B. Jussieu, not Plumier as the source of the collected material.In examining , I found numerous geographical notes provided by Linnaeus that, in light of our present understanding, seem to misrepresent the actual ranges of the species to which these notes refer. A selection of extant species published in 
bearing the identical geographical note (Habitat in America meridionali.) by Linnaeus yields Acalypha australis [known range, East Asia], 
Ruellia (Barleria) coccinea [known range Caribbean], [known range, pan tropical]. Linnaeus™s other 1753 
Iva species, Iva frutescens carried the geographic note: 57
. There are no known 
Iva species native to South America (Jackson 1960). 
The known present range of Iva frutescens L. is almost entirely coastal, from south Texas to Maine 
and Nova Scotia possibly excluding New Brunswick (Jackson 1960).In 1803, Andreas Michaux included 
Iva in his Flora Boreali-Americana
 (Michaux, 1803). He -cies in the Midwest. His description of the genus Iva, and that for Iva annua are shown below (trans-I V A   L.
Masc. (staminate), Calyx communis (involucre) 5-leaved, Corolla, single petaled, 
long. Seed, obtuse.ANNUA.  L. I. annual, hirsute: leaves oval-lanceolate, sparsely serrate: spike crowded; bracts acuminate, bracts and involucre hirsute.OBS.  Leaves opposite, then alternate, like in . Spikes almost like 

HAB.  Habitat in the Illinois area.How Michaux connected the Illinois specimens with Linnaeus™s description of 
Iva annua, is un-58
Iva annua L. is given separate status from Iva annua Mich. as for instance in Rydberg™s North American Flora (Rydberg 1922) where the Michaux version is seen 
as synonymous to the later designation Iva ciliata,  synonymous to Iva annua of Linnaeus. In 1804, all this confusion became much more permanent when Karl Ludwig Willdenow (1804) 
described Iva ciliata in his 
Volume III, part 3, page 2386.
 Al-though since Jackson™s work (Jackson 1960) we know 
I. ciliata as the synonym of I. annua, he thought, at the time that I. annua sensu Michaux was a distinct species from I. annua sensu Lin-naeus. The substantive difference in the species descriptions are the geographical notes at the end of 
each; namely for 
and for 

Willdenow cites Linnaeus; 
Hortus Upsaliensis, and Amoenitates Academicae 3, 
page 25; but also cites the illustrative plate, number 16 of Iva annua from Schmidel™s 
-rum (Schmidel 1762, 1793). The original  was released in 1762. The 
representation here (Figure 3-4), was taken from second edition (1793). I believe they are identical. A critical difference between the anatomical details depicted in Schmidel™s illus
-tration and wild Iva annua, is that the style lobes in the illustration are shown with 2, 3, and 4 lobes, in contrast to the 2 lobes seen in the wild and noted in Hortus Upsaliensis (Linnaeus 1748). Recall that Linnaeus stated he was working with greenhouse-grown specimens. Jackson (1960, page 808) states that when he grew Iva annua in a greenhouse over the winter, he produced specimens with 
variable number of style lobes as well, though ascribing this effect more to day length considerations 
than to cultivation per se. Willdenow™s reference to Schmidel™s plate is the earliest I have found, so 
59it appears as though Schmidel™s plate is thought to support Willdenow™s separate diagnosis for 
I. an-nua and I. ciliata. Linnaeus clearly states (above) in his 1748 (Hortus Upsaliensis) that the pistillate 
he adds underneath Iva ciliata (translation from Latin, and with parenthetical terms added by me). fiJust 
like the previous (Iva annua), but it seems to differ by: thick ciliate petioles, bracts ovate-lanceolate, 
sharp-tipped with very long cilia (hairs).fl
-tions, the crux of the matter seems to be the incorrect interpretation about South American distri
-bution from the original description by Linnaeus.  From Willdenow™s 1804 publication until R. C. 
Jackson™s 1960 reorganization of the genus 
Iva (Jackson, 1960), Iva ciliata stood, and Iva annua fell into disuse as an uncertain species.In 1836, of a variety, 
Iva ciliata var.
 latifolia, was added by de Candolle (de Candolle 1836 page 529). He credited the receipt of his material to his former student Jean Lois Berlandier, who was 
chosen by de Candolle to make botanical collections in Mexico, starting in late 1826. DeCandolle™s 
description of Iva ciliata latifolia is below (translation by me.) with three main veins and having widely separate teeth, alternate and lanceolate in the upper parts, bases of the petioles ciliate, bracts oval with very sharp tips, lobes of the involucre rounded with ciliate trichomes. Collected in Mexico, near Bejar by Berlandier. Is this a different species than the one from Illinois?fl
The collection location cited by De Candolle, Bejar, also known as Béxar, Mexico, was the site of 
a fort and small village that is now part of modern-day San Antonio (in Bexar County), Texas. An 
image of an isotype is shown (Figure 3-5). Although it has condition issues, it is clearly not so broad-
60Figure 3-4 
 Schmidel™s in˜uential plate of 1793 that may have confused Willdenow 

Icones Plantarum, published in 1793, showing the details of Iva annua 
Species Plan-tarum. 

Iva annua sensu Michaux was a different species than 
Iva annua sensu Linnaeus.61leaved as to stand out from the majority of I. ciliata (now I. annua) in herbaria across the United States. Iva ciliata var. 
latifolia by de Candolle appears to have been generally recognized as unre-markable and was included in Iva annua (I. ciliata at that time), from then forward.In 1899, John K. Small (1899) published Iva caudata from the swamps of Louisiana and Missis-sippi. He wrote (1899, page 290), fiIt may be distinguished from Iva ciliata by the smoother foliage, epithet, caudata, translates to elongated linear bracts, its most compelling feature as revealed in the type specimen shown (Figure 3-6).Small™s description of 
Iva caudata was foreshadowed in 1835, in a description by John Torrey, 
in the Companion to Botany Magazine (Torrey ex Hook 1835, pages 99-100) where he designates 

Although Iva annua, the text details two varieties (translation of the Latin portion by me, following the original write-up). 545. 
Torr. MSS.; hirsuto-scabra, foliis ovato acuminatis 
subinciso-serratus, racemis paniculatis capitulis longe bracteatis Š bracteis ovata acuminatis valde ciliatus.Š. bracteis lanceolatis vix ciliatus Š.N. Orl. 1833.[Rough hairy, leaves ovate and pointed with somewhat incised serrations, panicu
-late racemes of capitula with long bractsŒ bracts ovate, pointed and very ciliate. . bracts lanceolate and scarcely ciliate. . New Orleans.1833]ŠA most distinct plant, differing from the original 
(from the Red River) of Dr. Torrey in my Herbarium, in the somewhat narrower leaves and much 

-tinctly ciliated. The presence of these large bracteas readily distinguishes the species.
 545. 
Torrey certainly seems to be pointing at two ‚morphs,™ 
 and  of this species, recognizable by the 62Figure 3-5 
 The syntype of 
Iva ciliata 
variety
 latifolia 
de Candolle 
Iva ciliata variety latifolia de Candolle. Although it has condition issues it is 

Iva ciliata Iva annua-

63
 The 
Iva caudata
 syntype 
 Iva caudata syntype. This is one of the four type sheets des-64
Iva, not an 
, this name is listed as a synonym of Iva annua. Asa Gray, however took note of this in the 
Synoptical Flora of North America, 
(Gray, 1886
, page 246), where he stated, under I. ciliata, fi
Torr. in Hook. Comp. Bot. Mag. i. 99, 
with a var. having linear and much elongated bracts to the spike.fl 

-tify the descriptions of the, ficonspicuously elongated linear bractsfl  (Small 1899, page 290) in his text. In Blake™s paper (Blake 1939), in which he establishes 
Iva ciliata var. 
macrocarpa
, he ends the work with a long paragraph (pages 85-86) in which he makes the case for recognizing the species, Iva caudata, as a synonym of Iva ciliata .fiA form of the 
Iva ciliata group has been described by Small as Iva caudata, and is maintained by Rydberg in the fiNorth American Flora.fl The only really distinctive feature that appears in their keys 

fiovate to lanceolate, short-acuminate, hispid-ciliate along nearly the whole marginfl in 
I. ciliata, and filinear or linear-lanceolate, caudate-acuminate, ciliate only at the basefl in 
I. caudata. Small™s key 
character is the same, except that he does not mention a difference in the pubescence of the bracts. 
Iva caudata was originally described from Louisiana and Mississippi, but the range of the two as given by Rydberg is essentially the same, except that 
I. caudata is given a range from Illinois and Missouri to Mississippi and Louisiana, while I. ciliata is permitted to grow from the same eastern limit west to Nebraska and New Mexico. Examination of the specimens in the United States National 
Herbarium shows that the attempted separation corresponds to nothing in nature. The bracts vary from 


-etal distinction can be drawn. The alleged difference in pubescence of the bracts mentioned by Rydberg 
is non-existent. In his original description Small stated that the leaves were thinner and smoother. This 
is obviously an ecological feature associated with growth in a damp, shady habitat. A specimen from 
Mississippi labeled I. caudata by Rydberg has relatively thick, rough leaves, as do others from Texas 
Iva caudata by 
Rydberg. Iva caudata must be referred outright to the synonymy of 
I. ciliata.fl65This (Blake 1939) seems to be virtually the last time that Small™s (1899) characteristics of thinner 
smoother leaves are recommended as distinguishing features between I. ciliata and I. caudata. How-ever, the length-to-width appearance of the bracts, in accordance with Blake™s reminder, fi
extremes 
are naturally quite different in appearancefl 
Archaeological Discovery
In 1924, M. R. Harrington (1924), in an article, describes the discov-ery of shelters, natural features of cliffs derived from sedimentary layers in the state of Arkansas. In 
these dry, well-preserved sites, the remains of a culture(s), seeming to be pre-colonial were recovered 
in abundance. Within what were described as seed bags, preserved by the dryness, were what Har
-
-cally as Iva. Iva xanthifolia, although 
Iva ciliata. 
on page 101; fiA very interesting and curious fact is that the seeds in the stores were of a size much 
larger than any now growing as weeds. This suggests that all these larger seeds in the stores of the 
Bluff-Dwellers may have been the product of cultivation. The purpose for which is problematic.fl
Gilmore describes gathering some Iva xanthifolia seeds and being pleasantly impressed with the aroma and suggests that they may have been a perfume product (Gilmore 1931, page 87). But the Iva annua fruits, cypselae [achenes from an inferior ovary may be termed cypselae (Marzinek et al. 2008)] as the dominant component in some human paleofeces found in the 66cave systems of the mid-United States, and the nutritional revelations, including kernel composition of 32.25 percent protein and 44.47 percent fat, published by Asch and Asch (1978) make a compel
-ling case for their development and use as a food source (Wagner & Carrington 2014).
In 1939, Blake describes the variety Iva ciliata var. 
macrocarpa, 
to be comprised of the archaeo-logical specimens of such a large size that, fimakes it desirable to differentiate them by a varietal 
name, even though it is most probable that they represent merely an ancient cultivated strain obtained 
by selection, and now extinctfl (Figure 3-7, in comparison to wild seeds from the present, Figure 3-8).  In Blake™s description establishing the variety 
I. a. var. 
macrocarpa, 
of the cultivated-type cypselae as measuring 4.8 to 9.3 mm in length, and 3.2 to 5.7 mm in width (Blake 1939). Presently however, cypsela length measurements of over 4.0 mm are considered to 
have been under cultivation (Wagner & Carrington 2014, page 74).
Blake™s description of the large cypselae of the new variety complete his analysis, except for 
measurements of two phyllaries found in one of the bottles of cypselae. They are described as differ
-ing by size only, from analogous anatomy in modern 
Iva ciliata ( now I see this varietal distinction as worthwhile, because of its exclusively archaeological origins. In the same way that e.g. I. asperifolia var. 
asperifolia, I. asperifolia var. 
angustifolia, and I. asperifolia var. 
latior); the tem-poral range of I. a. macrocarpa 
supports its incorporation into the taxonomy of I. annua.In 1960, R. C. Jackson published A Revision of the Genus
 Iva L. In this thorough work, he estab-lishes the name Iva annua L. in priority over the long-established synonym Iva ciliata Willd. On 
page 808, he re-asserts the original name from Linnaeus and explains the problems with Schmidel™s 
67plate (Schmidel 1762, 1793) and how the abnormal presentation of day length, not simply green-house cultivation per se, is probably implicated in the non-typical number of stigmatic lobes as well as diagnosing the fiAmerica meridionalifl problem in the original description (Linnaeus 1753).Iva ciliata var. 
macrocarpa 
as a synonym of Iva annua var. 
mac-rocarpa, 
a comb. nov. for the large-fruited specimens from archaeological discovery; mostly repro
-ducing the diagnostic details provided in Blake™s original designation
.Jackson also maintained the varietal standing of what has now become Iva annua var. 
caudata comb. nov., replacing 
Iva ciliata var. 
caudata. In his description of Iva annua var. 
caudata, he speci-cauda-ta, leaves 
above. On page 855 (Jackson 1960) he shows two line-cut illustrations to demonstrate the intended contrast in appearances (Figure 3-9).Iva annua var. 
annua DiagnosisAnnuals, (10-) 50-100 (-150+) cm; Stems: erect with a short taproot, having mostly opposite lower branches. Leaves: petiolate, petioles 5-30 mm, blades deltate or ovate to elliptic, trullate or lanceo-late, 30-150 mm by 10-95 mm, margins with variable, widely spaced rounded teeth, surfaces scabrel
-to gradually become alternate. Heads: in axillary and terminal spiciform arrays, each subtended and 68exceeded by a bract. Bracts ovate to broadly lanceolate to slender caudate, ciliate-margined. 
Pedun-cles: 1-3 mm.  Involucres:
 mostly hemispheric, 3-5 mm. Phyllaries: 3-5 distinct and herbaceous. Paleae: linear, 2-2.5 mm. 

3 (5), corollas 5-lobed, 0.5-1.0 mm. 
8-15; corollas 2-2.5 mm, anthers and pollen pale yellow. 
Cypselae:-
 Cypselae from 
Iva ciliata
 (now 
annua
) var. 
macrocarpa 

Iva ciliata var. 
macrocarpa

-

 Cypselae size spectrum from Little Salt Fork, Lancaster Co., Nebraska 
Iva annua 
69axially concave/convex, 2.0-4.0 (6.0) mm. Smaller sizes rounded and smooth on both sides, larger 
size extremes 3-ribbed on the concave surface and up to 5-ribbed on the convex surface; brown to very dark gray, often dotted with amber colored resin dots. Irregularly common on the shores of the 
Mississippi and its tributaries and on disturbed sites throughout. Appears to reach large plant size on 
Mississippian sites (115+ cm), and maintain a smaller maximum size in saline conditions and alka
-When these plants perish and dry in late summer and fall, these sharp-tipped hairs become stiff and 
glassy, and can penetrate the skin of those handling plants or harvesting cypselae. While most plants 
start out green, many, during fruit set, especially in Mississippian sites become very darkly pigment
-ed from reddish to blackish purple, either on stems, or throughout. As indigenous Native Americans perfected their foraging, large stands of riverine weeds that 
produced plentiful seed would have attracted their attention. As often as they faced starvation 
(Diamond 2005) and as they certainly exploited every resource at hand, they would not have passed 
 Jackson™s (1960) depiction of the de˚ning characteristic  of 
I. annua 
var. 
caudata 

Iva annua. 
Iva annua var. 
annua, and 
Iva annua var. 
caudata.
-70up a nutritious seed resource (Asch and Asch 1978). The fact that this species was a stand-forming 

waterfowl, lends this plant to numerous encounters and familiarity. The fact that they brought it not 
variety we associate with their early agriculture, and that after a minimum of three millennia they let 
to the changing fortunes of the Native American relationship to agriculture and survival. 
ConclusionThe taxonomic history of the Asteraceae species known today as 
Iva annua, is one of the more convoluted stories in plant natural history. R. C. Jackson in his revision of the genus 
Iva (Jackson 1960) recognized Iva annua, the Linnaean species of 1753 from the mid-continent of North America, 
as the senior synonym of Iva ciliata Willd. Of the three varieties maintained by R. C. Jackson (1960) 
one of them, I. a. var. 
caudata, should be regarded as a junior synonym of I. a. var. 
annua, see Chap-ter 4. The taxonomy of genus Iva has gone from using a few scant characters, to detailed morpho-regions of chromosomes.71REFERENCES72REFERENCESASCH, D. L., N.B. ASCH. 1978. The economic potential of 
Iva annua and its prehistoric importance   in the lower Illinois Valley. 

 Michigan 67:300-341.AUSTIN, D. F. 2004. 
. Boca Raton, CRC Press.
BLAKE, S.F. 1939. A new variety of 
Iva ciliata from Indian rock shelters in the South-Central   
 United States. Rhodora 41:81-86.CORRELL, D. S., CORRELL, H. B. 1982. 
 Caicos Islands. Hirschberg, Germany, Strauss and Cramer.
DE CANDOLLE, A.P. 1836. 

. Paris.DIAMOND, J. (2005). The Ancient Ones: The Anasazi and Their Neighbors. 
Collapse: How Societies   Choose to Fail or Succeed. New York, Viking: 136-156.
DIGGS, G. M. J., LIPSCOMB, B. L., O™KENNON, ROBERT J. 1999. Shinner™s and Mahler™s Illustrated 
  Flora of North Central Texas
. Fort Worth, Texas, Botanical Research Institute of Texas.
FERNALD, M. L. 1950. Gray™s Manual of Botany
. New York, American Book Company.
GILMORE, M.R. 1931. Vegetal remains of the Ozark Bluff-Dweller culture. 
 Academy of Science, Arts and letters
. 14:83-102.GORTO
N, J. 1847. 
. London, Whittaker and Co.
GRAY, A. 1886. Iva. 

. 1:245-247.HARRINGTO
N, M.R. 1924. The Ozark Bluff-Dwellers. 
American Anthropol
ogist 26:1-21.HICKMAN, J. C. 1993. 
 Berkeley, University of 
  
 California Press.JACKSON, R.C. 1960. A revision of the genus 
Iva L. University of Kansas Science Bulletin  41:793-876.JARVIS, C. 2007. 
. London, The Linnean 
  Society of London.LINNAEUS, C. 1737. Hortus Cliffortianus. Amsterdam.73 1748. Hortus Upsaliensis. Stockholm, Laurentii Salvii. 1753. . Uppsala, Laurentii Salvii. 1754. . Stockholm, Laurentii Salvii. 1756. . Amoenitates Academicae 3:1-27.
LINNAEUS, C., CHENON, L.J. 1751. 
. Upsalla,  Academae Upsaliensis.MARZINEK, J., O.C. DE-PAULA, & D.M.T. OLIVEIRA by considering anatomical and historical factors. 
. 31: 1-6.MATASYOH, J. C., J.J. KIPLIMO, N.M. KARUBIU, T.P. 
HAILSTO
RKS. 2007. Chemical composition and   antimicrobial activity of essential oil of Tarchonanthus camphoratus.
 Food Chemistry   
 101:1183-1187.
MICHAUX, A. 1803. 
Flora Boreali-Americana
. Paris.NUTTALL, T. 1818. 

.   Philadelphia, D. Heartt.RYDBERG, P.A. 1922. Tribe 1. Iveae. 
North American Flora
. New York, New York Botanical 
 Garden. 33: 3-13.SAFFORD, W.E. 1924. Letter to M. R. Gilmore, April 17, 1924, concerning specimens of Ozark 
  
 Bluff-Dweller material collected by M.R. Harrington. M.R. Gilmore. Ann Arbor, Michigan, 
  Unpublished.SCHMIDEL, C.C.  1762. I Erlangen, Ioanne Christophoro Keller.
 1793. Erlangen, Johann Jacob Palm.SMALL, J. K. 1899. Undescribed species from the southern United States. 


 Gard.
 1:278-290.SMITH, B.D. 1992. Rivers of Change. Washington, Smithsonian Institution Press.
SPRAGUE, T. A. 1955.
 fiThe Plan of the Species Plantarum.fl 
   London 165: 151-156.STEARN, W. T. 1957. 




 London, The Ray Society. I: 1-176.
74STEARN, W. T. 1962. fiThe Origin of the Male and Female Symbols of Biology.fl 
 Taxon
 XI(4): 109-113.
THIJSSE, G., J. F. 
VELDKAMP. 2013. 
Leiden, Brill. Available at 
 www.brill.com/herbarium-van-royen-1704-1779. Accessed November 2014.
TORREY, J. ex HOOK, 1835. Ambrosia Pitcheri. 
Companion to Botany Magazine 1:99-100.TOURNEFORT, J. P. d. 1700. 

. Paris, Typographia Regia.
VAILLANT, S. 1719. De l™éablissement de nouveaux Caracteres de Plantes a Fleurs composées. 
  
 Classe II. des Corymbiferes. 

 1719:277-318.VAN ROYEN M.D., A. 1740. 
Florae Leydensis. Leiden, Samuelem Luchtmans.
WAGNER, G. E., P. 
H. CARRINGTO
N, 2014. Sumpweed or marshelder (
Iva annua).  In: Minnis P.E. ed. 
  New lives for ancient and extinct crops
. P. Minnis. Tucson, University of Arizona Press. 
  
 3:65-101.WEBER, W. A. 1987. 
Colorado Flora Western Slope.
 Boulder, Colorado, Colorado Associated 
 University Press.WILLDENOW, C.L. 1804. IVA. 
Linnaei species plantarum. Berlin. III, pt. 3:2386-2387.75CHAPTER 4A revision in the taxonomy of 
Iva annua to place the variety Iva annua caudata in synonymy with I. a. annuaIn 1899, when John K. Small (1899) published his description of Iva caudata collected from the swamps of Louisiana and Mississippi, he wrote that it may be distinguished from Iva ciliata by the -rescence. The epithet, 
caudata, translates to character of the elongated linear bracts, its most compelling feature as revealed in the type descrip-tion. He designated 4 type specimens, on deposit at the Columbia University Herbarium (one of which is displayed as Figure 3-6)

Certainly Blake™s (Blake 1939, pages 85-86) discussion of Small™s 1899 description shows the scepticism 
with which he regarded Small™s 
I. caudata designation. 

-rowness of the bracts, there is no other claim upon which to make a determination to identify I. a. var. 
caudata. Although Blake claimed that, fiExamination of the specimens in the United States 
National Herbarium shows that the attempted separation (into varieties) corresponds to nothing in nature,fl he did not actually do any measurements from herbarium specimens to solidify that claim.In Jackson™s Revision of the Genus 
Iva (Jackson 1960), by restoring Iva annua L., and making Iva 76ciliata a junior synonym of I. annua, he creates the new combination Iva annua var. 
caudata (Small) Jackson; seemingly accepting the validity of varietal status for Small™s former species (Jackson 1960).
much as provides a thorough review, and for the critical feature, namely the measurements of the 
Jackson, in his description of Iva annua var. 
caudata, describes, the remaining character for diagnosing the assignment of the name var. 
caudata, namely (see Figure 3-9) that he includes under the heading of leaves 
lanceolate 7-18 mm. long, caudate-acuminate, hispid-ciliate;fl but including no corresponding width I hypothesized that there is a smooth continuum of variation in the length to width ratios of the Iva annua. Since there is complete overlap between the ranges of I. a. cau-data and I. a. annua, a discovery that there is a smooth continuum between the two ‚morphs™ would 
I. a. var. 
caudata in synonymy with I. a. var. 
annua.Materials and MethodsIva annua, I have measured the lengths and widths ratios of approximately 190 Iva annua specimens from her-barium specimens graciously loaned to me by the University of Florida, the University of Texas, and 
Louisiana State University herbaria using the following procedure. From each herbarium specimen 2 most proximal capitula as well. The 3 bracts were measured, length and width, with a Wild-Heer
-77brugg glass microscope stage reticle in combination with a Bausch and Lomb dissecting microscope (with the scale in increments of 0.1 mm), and recorded into an Excel spreadsheet. Each specimen™s 
mean bract-length and bract-width was used to calculate the mean bract-length/width ratio. In Figure 4-2, these ratios were plotted in order of magnitude, least to greatest, for comparison.Results and DiscussionThe charted results include four red lines that depict where the length to width ratios data for Small™s declared syntypes fall in the spectrum of measurements. Figure 4-1 shows clearly, in support 
of the contention of S. F. Blake (1939), that there appears to be no basis for selecting any portion of 
the smooth distribution of length to width ratios as indicating that any of the extremes of this shape/ratio deserve varietal status. There are also in Figure 4-1 two black dots connected by lines with the two illustrated sketches 
-tion of the two varieties fall in the measured variations. In Turner™s (2009) assignment of the varieties of 
Iva asperifolia, morphological variations support given varietal names as they are followed in their variation across the long, continuous range of the species. In Iva annua, the range of variation of the specimens designated by the characteristic elon-I. a. caudata, completely overlap the range of variety I. annua var. 
annua. I believe that when evaluated with the information in Figure 4-1, the two extremes of variation cannot be given varietal status.Iva annua 78comprised of all the extant specimens of the living populations, namely Iva annua var. 
annua, and second, the large-fruited, now apparently extinct product of early American Indian agriculture in the 
eastern United States, Iva annua var. 
macrocarpa.
ConclusionIn conclusion, I recommend that the subspecies, Iva annua var. 
caudata, be considered a synonym of Iva annua var. 
annua. Perhaps there should also be a taxonomic review of whether the large size 
020406080100120140160180Bract of Iva annua caudataRedrawn from R. C. Jackson, 1960L/W = 9.89Bract of Iva annua annuaRedrawn from R. C. Jackson, 1960L/W = 3.0605102015Figure 4-1 
 Bract Ratios of 
Iva annua
 Herbarium Specimens
 The length to width ratios of the in-
Iva annua. The four red-colored lines indicate measurements 

Iva caudata


-
Iva caudata
79of the cypselae of the cultivated variety Iva annua var.
 macrocarpa
 from archaeological discovery is 
Socorro Co., NMMandan, NDOakland Co., MIIva annua annuaIva annua caudataFigure 4-2 
 The range of the present varieties of 
Iva annua 
The range of the varieties Iva annua var annua and Iva annua var caudata. The range of I. a. caudata 
of I. a. annua. This is further evidence for the synonymy of I. a. caudata with I. a. annua, as there is no 


80REFERENCES81REFERENCESBLAKE, S. F.
 (1939). fiA New Variety of Iva Ciliata From Indian Rock Shelters in the South-
  
 Central United States.fl Rhodora 41(483): 81-86.JACKSON, R. C. (1960). fiA revision of the genus 
Iva L.fl University of Kansas Science  Bulletin 41: 793-876.SMALL, J. K. (1899). fiUndescribed Species from the Southern United States.fl 
Bulletin of the   

 1: 278-290.TURNER, B. L. (2009). fiTaxonomy of 
Iva angustifolia and I. asperifolia (Asteraceae).fl  91(1): 76-83.82CHAPTER 5Protein and Lipid Content and Composition of Modern Populations of 
Iva annuaIntroduction
The fact that Iva annua L. (marshelder, sumpweed) was not been generally recognized as a 
food plant until nearly the middle of the 20th Century implies there is not a long history of aware-ness or interest in it as a nutritional feature of the human diet. When 
Iva annua-
food plant (Gilmore 1931). It had not appeared in any of the chronicles of the useful plants of 
the American Indians written since European contact. It was variously suggested that it might 
have been a medicinal or even a perfume plant (Gilmore 1931). After the discovery of paleofeces 
from the Newt Cash Hollow (Jones 1931) and the nutritional assay from Asch and Asch (1978), 
and paleofeces from the Big Bone Cave (Faulkner 1991) and from the Mammoth/Flint Ridge 
Cave system (Gremillion and Sobolik 1996) that were high in consumed I. annua seeds, it was 
ultimately accepted as having been a food plant among the ancient American Indians of central 
North America. 

seeds of I. annua. In addition to discovering important vitamins and minerals (Table 5-1), their 
discovery of the high oil, high protein fraction of these cypselae propelled this species into no-tice. In Jared Diamond™s well-known essay, 
Guns, Germs, and Steel, he goes so far as to say I. 
annua, fiin particular, would have been a nutritionist™s ultimate dream, being 32 percent protein 
and 45 percent oil,fl (Diamond 1999, page 151).83Protein ŒThe importance of protein to the diets of all known vertebrate species has been a recog-nized feature of biology for over a century. When Asch and Asch (1978) published evidence showing 
that Iva annua seed had a protein content of 32.25 percent, it helped to end the debate about whether the seeds of this plant were used as a food. Asch and Asch (1978, page 303) with regard to their 
protein analysis reported; fiA sample of achenes was sent to Analytical Biochemistry Laboratories, 
Inc., Columbia Missouri, for determination of nutritional composition.fl The clear implication is that 
all their nutritional metrics were determined by the analysis of this single sample. Curiously, for the 
large interest generated by this reported protein level, no one has repeated these measurements.
Additionally, Asch and Asch calculated that their single sample had a protein content in proximate 
analysis of 32.35 percent using an N to Protein conversion factor of 5.30 (Asch and Asch 1978)  as 
reported on page 306.  This is consistent with D. B. Jones™s (Jones 1941) table (No. 5, page 14) 
wherein he reassessed the calculation of N to protein conversion factors and recommends the use of 5.30 x N for determination of protein in H. annuus seed. Asch and Asch took this to account as the 
proper conversion factor for oily seeds as it is the identical conversion factor Jones recommends for 
 Selected Mineral and Vitamin Comparison (mg/100g edible portion) of 
Iva annua 
to Two Modern Asteraceae Seed Crops
 
Iva annua, Marshelder*CalciumPhosphorusIron%DV%DV%DV%DV%DV%DV%DVPotassiumThiaminRibo˜avinNiacinVitamin B1Vitamin B2Vitamin B3290130011.47802.130.7513.1Helianthus annuus, Sun˜ower*1208377.19201.960.235.4Carthamus tinctorius, Sa˚ower77.86642.865685.711000mg1000mg18mg3500mg1.0710.3572.1431.5mg1.7mg20mgSelected Mineral and Vitamin Content (mg/100g edible portion) of Three Asteraceae Seed Crops* Numbers taken from:  Asch and Asch, 1978 (Table 3, page 307) Numbers taken from:  nutritiondataself.com/facts/nut-and-seed-products/3068/229%12%7.79%130%83.7%64.3%63.3%39.4%27.8%22.3%26.3%19.6%142%130.7%71.4%44.1%13.5%21%65.5%27%10.7%Recommended Daily Values˛ (www.dsld.nlm.nih.gov/dsld/dailyvalue.jsp)˛ Numbers taken from: www.dsld.nlm.nih.gov/dsld/dailyvalue.jspIva annua, Marshelder*CalciumPhosphorusIron%DV%DV%DV%DV%DV%DV%DVPotassiumThiaminRibo˜avinNiacinVitamin B1Vitamin B2Vitamin B3290130011.47802.130.7513.1Helianthus annuus, Sun˜ower*1208377.19201.960.235.4Carthamus tinctorius, Sa˚ower77.86642.865685.711000mg1000mg18mg3500mg1.0710.3572.1431.5mg1.7mg20mgSelected Mineral and Vitamin Content (mg/100g edible portion) of Three Asteraceae Seed Crops* Numbers taken from:  Asch and Asch, 1978 (Table 3, page 307) Numbers taken from:  nutritiondataself.com/facts/nut-and-seed-products/3068/229%12%7.79%130%83.7%64.3%63.3%39.4%27.8%22.3%26.3%19.6%142%130.7%71.4%44.1%13.5%21%65.5%27%10.7%Recommended Daily Values˛ (www.dsld.nlm.nih.gov/dsld/dailyvalue.jsp)˛ Numbers taken from: www.dsld.nlm.nih.gov/dsld/dailyvalue.jsp84hazelnut (Corylus avellana), walnut (
), coconut (Cocos nucifera), cottonseed (Gossy-pium hirsutumLinum usitatissimum) (Jones 1941).Initially, when most known proteins were of animal origin, it was discovered that most proteins 
then known were comprised of an average of 16 percent nitrogen. This lead to the simple calculation 
(100 / 16 = 6.25) that 6.25 is the conversion factor appropriate for determining the protein content of 
it assumes that all nitrogen is representative of protein in the sample. Of course this ignores the free amino acid content and peptides that may contribute to the measurement of percent N. It seems, in the face of these considerations, this style of conversion should properly be considered a measure-ment of total included amino acid, in preference to available protein per se. In the FAO/WHO guide, 
Protein Quality Evaluation (FAO/WHO 1991, page 20) a recognition is made that since inter-labora
-tory reproducibility of amino acid analysis numbers is within about 10 percent (Section 5.5 Conclu-sions and Recommendations, point 1.) that further work be devoted to improving the accuracy and reproducibility of the analysis procedures, and that (point 4.), fiAmino acid data should be reported as mg amino acid/g N or converted to mg amino acid/g protein by use of the factor 6.25. No other 
modern literature although its problems are increasingly recognized (FAO/WHO 1991).
D. B. Jones (1941) however, made clear the pitfalls of using this universal conversion factor, 
based largely on the increasing knowledge of how differing compositions of amino acids with their 
variation in N content (from 1 N atom, in 14 proteogenic amino acids, to 4 atoms of N in arginine). 
For the example cited in his text he (Jones 1941, page 7) evaluates, for almond, the results of using 85the 6.25 x standard, against the results recognizing the actual N of almond protein of 19.3 percent (translating to a factor of 5.18). The factor, when tailored to the almond protein nitrogen level, yields 
a protein level of 17.4 percent instead of 21 percent, a notable difference.
both the following:Where  kAresidues per 100g of seed dry weight, over the sum of the grams of nitrogen recovered from the amino acids per 100g of seed dry weight. In comparison, kpbased upon the sum of recovered amino acid residues per 100g of seed dry weight, over the sum of the grams of total of nitrogen, measured as per the Kjeldahl method (Kjeldahl 1883). Mossé reports that, fithese two factors are the upper and lower limits, respectively, of the total seed N to true protein 
conversion factor k, which is close to the average of kA and kppage 23) that fiThe present results show that k varies from 5.13 for N-poor rice samples to about 6.0 for N-rich foxtail millet (Setaria italica) samples.flWorking backward from Asch and Asch™s report of 32.25 percent protein yields (32.25 / 5.30) a result of 
6.085 percent nitrogen for the kernels in their sample. Using the two k values as a bracket from Mossé™s 
work gives a revised protein content for the Asch and Asch sample as between 31.22 and 36.51 percent. 
The results of the C:N tests of samples from four populations (2 from freshwater origination, and 2 from 

k A  =  E i /  D i k p  =  E i /  N 86LipidsŒBesides the high levels of protein found in the analysis by Asch and Asch (1978), was the also 

of lipid, especially when appreciated concomitantly with the 32.25 percent protein level in the same sample. 
This probably contributed to why Jared Diamond considered sumpweed a ‚nutritionist™s dream™ (Diamond 
1997). Conspicuously absent from the nutritional assay of I. annua by Asch and Asch is an inventory of 
fatty acids comprising the abundant seed oil. As indicated (Asch and Asch 1978) a (single) sample, presum
-ably from the lower Illinois River Valley, was sent to the Analytical Bio Chemistry Laboratories, Inc., of 
Columbia, Missouri where the shells and kernels were each subjected to proximate analysis. Here Asch and 
Asch (1978, Table 1) report the fat content to be 44.47 g/100g of Iva kernels. This value was reported along 

I. annua in a 
Helianthus annuus
(Zea mays), as well as three species of wild tubers and four species of 
acorns and nuts, concluded their analysis of the fat or oil content of their Iva kernels.The present study revisited the study of Asch and Asch (1978) to more fully investigate the nutritive 
content of I. annua cypselae for their protein, amino acid, fatty acid and oil content and composition from 
wild populations representing both fresh water and saline environments.  I also tested the hypothesis that 
populations adapted to saline environments possess higher proline content and higher germination rates 
when tested under saline conditions compared with populations adapted to fresh water conditions.  This 
experiment was also an attempt to provide some insight into whether populations from the saline parts of 
the range have potential genetic adaptation to the saline environment that separates them in some sense 
from the non-saline inhabiting populations.87Materials and MethodsPlant MaterialŒThe samples of Iva annua cypselae were collected during this investigation, as shown in Table 5-2.  The cypselae noted in table 5-2 as, Heiser Site, Indiana were sent to me in 
late 2009 by the late Dr. Charles Heiser of Indiana University. His description of the locality was, 
fiIndiana University farmland south of Bloomington, Indiana.fl The Wagner Site seed was sent by 
Professor Gail Wagner (University of South Carolina) from plants she was growing near her home 
in Columbia, South Carolina. Plants were cultivated in the greenhouse during the 2011 and 2012 
seasons to produce seeds for the C:N assessment and some of the lipid compositional analysis. The 
seeds were soaked in deionized water and upon germination (1-15 days) transferred into 2.5 cm pots. After approximately 2 weeks in 7.5 cm pots, followed by approximately 6 weeks in 18 cm pots they were all transferred into 30.5 cm pots. Cypselae were harvested when the plants senesced in early 
 Collection Locations and Dates
 
collected Iva annua 
from farm acreage south of campus at Indiana University, Bloomington, Indiana.
’˚˛—fi˛fi˛ƒ‚˛˛‹’™fi˛fi˛fi˛fi˛fi˛fi˛fi˛ƒ‹‚˛˛‹’™fi˛ƒ‹‚˛˛‹’™fi˛–™˛–fi˛–fi˛–fi˛ƒƒƒƒƒ˜˛Œ˜˛Œ⁄˙˚ŠŸ88winter. They were all grown under supplemental lighting of 14 hours of light, 10 hours of dark, after 
Saline Wetland seeds in series c, Table 5-4) were not given supplemental fertilizers or salt.
Protein AnalysisŒIn order to establish the total nitrogen and total protein levels, samples were analyzed for C:N ratio and amino acid composition. The samples for the C:N determination were 
sent to the Duke Environmental Stable Isotope Laboratory at Duke University, Durham, NC. The 

 Parental Heritage of the C:N Ratio Samples
 
Iva annua cypselae from plants grown in the greenhouse from 


A1 PL-03 LSF  MD  5.56  4.78  26.64A2 PL-28 LSF LG  8.1  9.62  50.60A3 PL-27 III SM  4.79  4.52  61.29A4 PL-67 III LG  5.77  11.31  47.19A5 PL-25 III SM  12.0  9.07  48.84A6 PL-45 III MD  6.81  11.13  46.81A7 PL-37 III SM  9.57  9.40  48.60A8 PL-68 III LG  6.73  11.38  44.88A9 PL-08 PFZ MD  11.8  9.68  46.19B1 PL-27 III SM  7.13  0.71  44.04B2 PL-68 III LG  9.27  1.40  47.21B3 PL-45 III MD  9.76  1.13  49.20B4 PL-08 PFZ MD  8.02  1.79  46.92B5 PL-67 III LG  8.32  0.73  49.48B6 PL-25 III SM  10.19  1.26  43.60B7 PL-28 LSF LG  5.93  0.75  48.53B8 PL-37 III SM  12.67  1.15  48.64B9 PL-03 LSF MD  10.71  1.87  46.33PERICARPSKERNELSCultivated PltParent Sd SzSmp wt (mg)Sample% N% CParental Heritage of the C:N Ratio Samples89automated analyses of bulk carbon and nitrogen isotopes are performed on a Carlo Erba Elemental 
made (Table 5-2), there were 18 samples submitted, Table 5-3; 9 each of kernels and pericarps from 
populations representing both origins in freshwater and saline environments.  As the recommended sample weights (Will Cook, personal communication based upon the presumed 
N content) were between 5 and 10mg, the samples consisted of more pericarps than kernels. The parental 
heritage of the sample seeds are shown in Table 5-3; seeds for analysis were harvested from plants culti
-vated in the greenhouse during the 2011 and 2012 seasons. 
For the amino acid analysis, four samples of wild-collected seeds were submitted to the Chemical Lab-oratories of the University of Missouri-Columbia, College of Agriculture, Food and Natural Resources, 


The samples were sent including kernels with pericarps to be milled and tested at the Experiment Station 
Laboratories. There after milling, they were hydrolyzed and analyzed using cation-exchange chromatog
-
Lipid Analysis. 
marked on Table 5-2, see asterisks), 5 (5 kernels each) wild-collected from cypselae from popula
-tions growing in freshwater environments, and 4 samples (5 kernels each) from populations growing in saline environments were extracted as described below. Samples for the fatty acid compositional 
analysis, Table 5-4, utilized both wild-collected and cultivated (greenhouse grown) cypselae. Sam
-ples (1-5 kernels depending on which run) Table 5-4, were lyophilized for two days and weighed 
90extracted twice in 500 l of hexane/isopropanol (2:1). Combined supernatants were dried under a stream of N2. Lipid samples for fatty acid analysis were transmethylated with 0.5 ml methanolic HCl 
percent NaHSO4 in H2O were added and mixed vigorously. Fatty Acid Methyl Esters (FAMEs) were 
a Site 1  a Site 1  a Site 10 (L)  a Site 10 (L)  a Site 10  a Site 10  b Site 10 (L) b Site 10 (L) b Site 10 (L) b Site 10   b Site 10  b Site 10  c Site 10F  c Site 10F  c Site 10F  c Site 10  c Site 10  c Site 10  d Site 3  d Site 3  d Site 3 d Site 6  d Site 6 d Site 6  e Site 4  e Site 4  e Site 4 e Site 2  e Site 2  e Site 2  5.81mg 1 kernel 3.96mg 1 kernel 3.52 mg 1 kernel 3.03mg 1 kernel 3.31mg 1 kernel 3.17mg 1 kernel 2.93mg 1 kernel 4.05mg 1 kernel 1.49mg 1 kernel1.65mg 1 kernel 1.24mg 1 kernel 1.77mg 1 kernel 2.34mg 1 kernel 2.75mg 1 kernel 2.22mg 1 kernel 1.98mg 1 kernelf Site 5 f Site 5  f Site 5  f Site 8  f Site 8  f Site 8  f Site 11 f Site 11 f Site 11 g Site 5   g Site 9  g Site 5  g Site 5  g Site 5  g Site 5  g Site 9  1.43mg 1 kernel 1.50mg 1 kernel1.09mg 1 kernel  2.46mg 1 kernel 0.63mg 1 kernel 2.22mg 1 kernel 1.09mg 1 kernel 2.25mg 1 kernel 2.76mg 1 kernel 27.5mg 5 kernels  29.5mg 5 kernels 21.3mg 5 kernels 19.5mg 5 kernels 22.8mg 5 kernels 18.0mg 5 kernels 28.7mg 5 kernels g Site 10  g Site 5  29.3mg 5 kernels21.6mg 5 kernels2.19mg 1 kernel 1.87mg 1 kernel 1.14mg 1 kernel 1.14mg 1 kernel1.81mg 1 kernel 0.58mg 1 kernel1.53mg 1 kernel0.57mg 1 kernel 2.59mg 1 kernel  1.475mg 1 kernel 1.45mg 1 kernel 1.86mg 1 kernel 2.43mg 1 kernel 2.29mg 1 kernelThe Origin of the Samples for GC-FID AnalysisFreshwater PopulationsSeries     Site           Sample wt Kernels per *Site Numbers refer to Table 5-2  Site 10 (L) refers to cypselae from an individual (P˜zer) plant with exceptionally large fruits  Site 10F refers to plants that received three doses of fertilizer mid-summer (1 per week)  Miracle Grow® at 10ml/2.5 l water before ˚owering.Saline PopulationssampleSeries     Site           Sample wt Kernels per sample1.2.3.4.5.6.7.8.
9.10.11.12.13.14.15.16.17.18.19.20.21.22.
23.24.25.26.27.28.29.30.31.32.
33.34.35.36.37.38.39.40.41.42.43.44.45.46.
47.48.
The Origin of Samples for GC-FID Analysis 
The sources for cypselae samples used in 


received fertilizer.
91extracted into1ml hexane. Sample series a-g were analyzed with an Hewlett-Packard gas chromatograph, 

m id. 

I. annua lipid from 5 kernels from each of 9 populations (marked with asterisks in Table 5-2) was extracted as described above using benzyl ben
-
-
-ing the R-Statistics Package (R version 3.2.1 [2015-06-18] for Macintosh. The outliers were reported 
using the Bonferonni test parameters included in the R Statistical software package.Germination Tests. 
ΠIn order to contrast the germination success in fresh and saline conditions samples of wild-collected seed originating from both the fresh water and saline populations were germinated variously in 100% fresh, deionized water as well as 0.9 percent saline solution, and 1.8 percent saline solution in experiments conducted in both the summer of 2009, and 2013. In 2009 three samples (20 cypselae each) were germinated in each of the three salinity classes, from each of six populations (3 originating from freshwater populations and 3 from saline populations) for a total yield of 54 samples of 20 cypselae each (1054 cypselae). In 2013, the same procedure was used except that two, twenty cypselae samples were used instead of three (This was because seed stocks 92were running low on collected seed from the Mississippian and Nebraskan collections) for a total yield of 24 samples of 20 cypselae each (480 cypselae). Each Petri dish was lined with 90mm What
-
NaCl, 0.9 percent NaCl, or 1.8 percent w/v NaCl) then placed 27 cm below a 4 foot, 2 bulb high dark per day. Germinations were recorded each day until germination ended.
The 3 comparisons of performance of populations of fresh-origin to saline origin seed germination ANOVA using the R statistical analysis software for Macintosh, version 3.2.1 (Copyright 2015). 
ResultsTotal Protein. 
ŒThe percent C and percent N for each sample are presented in Table 5-3. For two 

implied by Bonferonni Outlier test), averaging 4.65 percent, or 45 percent of the average for the remaining samples, which averaged 10.23 percent. The lower values are similar to those of Asch and 
Asch (1978). For the other samples, the percent protein ranged from a low (sample A5) of 46.5-54.4, 
to a high (sample A8) of 58-68.3 percent protein using the two conversion factors discussed above. 
The average percent protein calculated for the higher N samples was 52.5 to 61.4. The average 
percent protein for the lower N samples (A1 and A3) was 23.9-27.9 and the average for all samples 
combined was 46.1-53.9Amino Acids. 
I. annua 
93Alanin
eArginine
As
xGlxGlycinePr
olineSerine
Histidin
eIsoleucineLeucineLy
sineCy
steineMethionine
PhenylalanineTy
rosineThreonine
Tr
yptophanValine05.015.025.010.020.0Amino AcidMass Percent of ProteinP˜zer Saline WetlandLincoln Saline WetlandAsch and Asch (fresh)Granite City, IL Site A (fresh)Wickli˚e, KY Site A (fresh)NonessentialEssentialFigure 5-1 
 Comparison of Mass Percent of Proteogenic Amino Acids by Freshwater and Sa
-line Sites 
The quantitative assay of amino acids from 
Iva annua 

˜˚˛˝˙˙˚˛˙ˆ˚˜ˇ˜˚˘



˙˚˛˝ˇ˚˙˜

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˘˚ˆˇ˙ˇ˚˛ˇˇ˚˜


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˙˚˛˙
ˇ˚˙˛
ˇ˚˝ˆ
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ˇ˚ˆˇ˘˚ˆˆ˘˚ˇˆ˘˚˝ˇ˚ˇ˘˜˚ˆ˝
˙˚˜˝
˘˚˘˛˙˙˚˛˙˝˚˝


˜˚˛˘
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‡‡



 Mass Percent of Amino Acids
 

Iva annua 

94three freshwater populations; namely the data from Asch and Asch 1978, and 2 collected from Gran
-ite City, Illinois, site A, and Wickliffe, Kentucky, site A. These three freshwater originating sites are 

-
Lincoln, Nebraska, are shown in shades of blue. These data establish the comparative percentages of 
the amino acid levels in each sample; in contrast to the C:N ratio measurements that become, with the consideration of the N to protein conversion factor, the better indication of the total bulk protein 
percent by weight of kernels from these sites. The amino acids that showed the greatest contrast from 
the freshwater populations to the saline populations were aspartic acid/asparagine, glutamic acid/ 
the analysis of replicate samples from each site, therefore no statistical inferences can be drawn 
different individual populations. However, when the three freshwater population samples are treated 
as replicates of freshwater plants, they can be compared with the two saline population samples. This 

Total Lipids.
 ŒAn annotated NMR spectrum of seed lipids is shown in Figure 5-2 (sample origi-nating in cypselae collected from Site II, Granite City, Illinois) depicting the contributions to the 



-95corporated into the TAGs in contrast to those that are free fatty acids. The contribution depicted from 
the anti-oxygen end of each fatty acid shows on the display whether or not the fatty acid is part of a TAG molecule. The signals (Figure 5-2, A) that permit the assay of the methyl groups at the ends of 

-gen atoms belonging to the glyceryl backbone at the ‚top™ of the TAG molecule (Figure 5-2, B) allow 
one to determine the TAG content. Since there are 3 component fatty acids in each TAG molecule, 

molecule, allow the calculation of the amount of free fatty acids from these cypselae.Graphic representation (Figure 5-3), shows the percentage of oil in the kernel samples from nine populations (5 from freshwater environments and 4 from saline environments). Figure 5-4 shows the Signal from these 9 H atoms in
the terminal (
) methyl groups
Signal from these 4 H atoms in
the glycerol backbone
Signal from these 6 H atoms next to

the carboxyl ends of the fatty acids (
)Triacylglycerol 

moleculeFigure 5-2 
 Sources of the main signals depicted in an NMR spectrograph 
Figure illustrates the 


I. annua cypselae.AB9650454035302520151050Percent Lipid of Cypselae by weightSaline SitesFreshwater SitesSite 1Site 2Site 5Site 6Site 4Site 9Site 8Site 10Site 11 TAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FAPercent Populations of Origin of Sampled CypselaeFigure 5-3
  Percent lipid in cypselae of populations from fresh and saline sites
 The gross 
percentages of oil as a percent of the weight from cypselae collected from 5 freshwater sites and 


1009080706050403020100PercentSite 1Site 2Site 5Site 6Site 4Site 9Site 8Site 10Site 11TAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FATAG   Fr FAPercent TAG vs Free Fatty AcidPopulations of Origin of Sampled CypselaeFigure 5-4 
 Percent TAG vs Free Fatty Acid
 Figure depicts the amount of fatty acids as TAG quanti
-

97Figure 5-5 
 Fatty Acid Pro˚les of Seeds from Saline and Freshwater Environments 
This graph 
Iva annua oil from cypselae from populations originating in freshwa
-
greenhouse environment under freshwater conditions.
Figure 5-6  
Box-plots of the 4 Iva Fatty Acids by Percent
  A 
B 
outlier. 
C 
test sample 22 is an outlier. 
D 
means. In a Bonferonni test sample 22 is an outlier.
ABCD01020304050607080
90C 16:0C 18:0C 18:1C 18:2Seed heritageSalt origin-wild collected   n=15Salt origin (cultivated in fresh water)  n=9Fresh origin-wild collected   n=15Fresh origin (cultivated in fresh water) n=6Fatty Acid Pro˜les of Seeds from Saline and Freshwater EnvironmentsNormalized Percent of Total Fatty Acid SD ± 1.80SD ± 1.42
SD ± 1.28
SD ± 0.98SD ± 1.55SD ± 0.52
SD ± 1.3SD ± 0.69SD ± 4.62SD ± 16.5
SD ± 5.85
SD ± 14.9SD ± 4.41SD ± 15.3SD ± 5.93
SD ± 14.616:018:018:118:298
and non-saline sites and for their progeny greenhouse-grown under fresh water cultivation. Figure 5-7 shows a typical display from the I. annua GC-FID results. Germination Tests.
 ŒIn the seed germination studies, the 0.0 percent salinity comparison, Figure 5-8 A50 percent of the total germinations on day 3 for cypselae originating from the saline populations and day 4 for cypselae originating from freshwater populations. The saline originating populations 
ultimately had approximately 5.83 percent greater possible germination success overall. In a lump-sum one-way ANOVA the mean score for the 5 freshwater populations was 36.59 ±11.32, and for 

 A characteristic result from the GC-FID analysis process
 This is typical of the results 


during the sample preparation.
16:017:018:118:218:099
Accumulated Percent Germination at Three Salinities
 A The germination record com
-

 B The germination record 


C The germination 


ABC100
p=0.0131. At 0.0 percent salinity, the success rate for saline originating seed exceeded the freshwater 
by about 13 percent.In the 0.9 percent salinity comparison, Figure 5-8 B, saline water originating population seeds achieved 50 percent of the total germinations on day 5, while the freshwater populations did not achieve 50 percent until day 11. Saline originating populations ultimately had ~13 percent higher 
germination success overall. In a lump-sum one-way ANOVA the mean score for all 5 of the fresh
-water populations was 13.94 ±7.14, and for the 5 saline populations 27.98 ± 10.27 (Average ±SD) 
and the difference at p= 0.001. At 0.9 percent salinity, the success rate for saline seed exceeded the 
freshwater by 13 percent.In the 1.8 percent salinity comparison, Figure 5-8 C, saline water originating population seeds achieved 50 percent of the total germinations on day 9, while the freshwater originating populations achieved 50 percent on day 14. Saline ultimately had approximately 9.44 percent greater germina-tion success overall, although this success rate was in a lump-sum one-way ANOVA the mean score 
for the freshwater populations was 1.27  ±0.82, and for the saline 7.21 ± 3.57 (Average ±SD) and the 

DiscussionTotal Protein. 
ŒIn Duke and Atchley™s Handbook of Proximate Analysis Tables of Higher Plants 
(Duke and Atchley 1986), in table 3 - Data converted to a zero-moisture basis, compiles a list of 
7342 analyses (not every number stands for a separate species; some species are listed for more than 101one analysis, and some species have several varieties on the list). Of the 32 species whose seeds are listed as being comprised of 50 percent protein content or greater, none are established crops. 
Duke and Atchley used published values; some used the conversion factor of N x 6.25, others were 
undisclosed. Since this 1986 compilation, high protein soy varieties, Glycine max, exceeding the 50 percent protein level have been achieved (Leffel 1992). Of the 8 species whose seeds were found to 
contain protein levels of 61 percent or more, only three are now known to have edible seeds (-
). Duke and Atchley report numbers 
for 
that average (n=5) 29.04 percent with 64.4 percent as the highest. The number for 
Heterophragma adenophyllum 
of 62.5 was the only report for that species. The reports for 

(n=9) average 30.53 percent with the highest being 65.2 percent. The majority of samples 
here exceeding 60 percent makes these among the highest protein levels of any plant species and the highest of any food plant cultivated by Native Americans or other indigenous peoples.
Amino Acids. ŒAsch and Asch (1978), strongly argue that while the protein content is high, the 
(page 303), when compared with the FAO 1973 reference pattern. It should be noted that when Asch 
and Asch (1978) was published, the FAO/WHO  1973 guidelines for establishing the suggested refer
-ence pattern for amino acids was based upon, fiuse of a single reference pattern to be applied for all needed some 30 percent of their protein in the form of IAA (indispensable [essential] amino acids) 
while the adult apparently needed only 15 percent or less.fl Since then, lowered values (FAO/WHO 
1991, page 21) have been adopted, that converts to higher amino acid scores for all upper age groups. 102
both aspartic acid and asparagine, and Glx represents both glutamic acid and glutamine) repre-sented in the results, 8 show higher levels throughout the freshwater or saline populations. The 
amino acids that showed the greatest apparent contrast from the freshwater populations to the 
saline populations were aspartic acid/asparagine, glutamic acid/ glutamine, arginine, and proline 

levels in plants dealing with abiotic stress, especially cold, drought, and salt tolerance. Proline 
as a soluble osmolyte is involved with mitigating damage from NaCl exposure (Xiong and Zhu 
2002). The modestly higher levels of proline in the seeds of the salt-originating populations 
indicates that I. annua may accumulate some free proline in response to saline conditions. Proline 

Helianthus annuus, seedlings exposed to moderate salt 
stress has been reported (Shi and Sheng 2005) and is common in salt tolerant plants. Salt stress 

-ance section of the Arabidopsis Book with fi...no toxic substance restricts plant growth more than 
does salt on a worldwide scale,fl (Xiong and Zhu 2002, page 1). When one encounters 
I. annua in 
-lations, from 0.6 to 1 meter (2 to 3 feet), instead of the 1.5 to 2 meters (4.5 to 6 feet), (Kaul 2006 
and personal observation).  When grown in the greenhouse without added salt, no dramatic differ
-ences were observed between the height of mature plants from saltwater and freshwater popula-
from single populations should be analyzed. To determine whether proline is accumulated as a 
103
especially roots, should be measured from plants grown under freshwater or saline conditions.Since the United States Department of Agriculture (Food and Nutrition Board, Institute of 


I. 
annua
Iva kernels 

RDA for lysine and threonine respectively. The idea that one (adult) could get the whole day™s protein 

plant origin.
  Iva annua
 kernel quantities required to meet USDA minimum levels of essential 
amino acids  
amino acids to the amount of Iva annua 
dietary minimums for these nutrients.       1USDA AA     
2Lowest Content    g 
Iva
 protein     g 
Iva
 kernels to 
   Requirement mg/day for  g/day for in 
Iva
 Protein  to meet RDA          meet RDA if 
Iva
    (mg/Kg/day) 70Kg body 70Kg body (g AA/g protein)       is 50% prot. 
Histidine    15 1050  1.05  0.0250   41.9     83.9
 Isoleucine    21 1470  1.47  0.0387   38.0     76.1 
 Leucine    47 3290  3.29  0.0661   49.8     99.6
 Lysine     43 3010  3.01  0.0305   98.8   197.6
 Methionine + cysteine  21 1470  1.47  0.0342   42.9     85.9 
 Phenylalanine + tyrosine  38 2660  2.66  0.0613   43.4     86.8
 Threonine    22 1540  1.54  0.0213   72.3   144.6 
 Tryptophan    6 420  0.42  0.0290   14.5     29.0
 Valine     27 1890  1.89  0.0519   36.4     72.8 
1Taken from USDA (Food and Nutrition Board, Institute of Medicine 2005).
2Lowest values for each of these AA (see Figure 5-4) taken from the 4 populations assayed by Carrington.
3             Amino acids with the lowest percentage of daily amino acid requirement per gram of kernel.
Iva annua
 kernel quantities required to meet
USDA minimum levels of essential amino acids
33104LipidsŒAs shown in Figure 5-9, the component fatty acids in the cypselae oils of I. annua, are not 
H. annuus and C. tinctorius

last much longer in intact seed than once pressed out. There is also an effect on vegetable oils of the 

at low concentrations, they lose that effect at high concentrations. Natural vegetable oils often seem 
to have  tocopherol concentration very near the optimum for the stability of their oils (Kamal-Eldin 2006). This investigation did not assay tocopherols for in 
I. annua. 
acid composition between saltwater and fresh water populations nor did cultivation of plants from saltwater under fresh water result in notable differences. For future work, the small sample size and 
the potential for high throughput make FAME analysis a potentially useful tool for assessing the di
-growth under controlled conditions.Asch and Asch (1978) originally reported 44 percent fat (oils) in their sample of 
I. annua. Analysis in this investigation found 22-28 percent lipid by weight, but 3 of my samples, see Figures 5-3 and 5-4, which have in addition to typical TAG levels, high levels of free fatty acids. The work by Asch 
105and Asch showed no evidence of any attempt to distinguish triacylglycerides (TAG) from free fatty 
acids. A possible factor in the levels of free fatty acids is storage time. Asch and Asch gave no clues 
to the interval between collection and analysis in their 1978 paper. In this study, storage times varied 
from four to seven years, during which time all samples were kept between 3 and 7 degrees Celsius. Storage times have been documented to have an effect on free fatty acid formation in cottonseed 
(Karon and Altshul 1944).
Also there is some evidence (Claassen, et al. 1950) that seed protein levels are inversely related to oil 
levels. The implication is that if Asch and Asch (1978) used a sample from a nitrogen poor environment, 
˜˚˜˛˜
˝˜
˙˜ˆ˜ˇ˜
˘˜n-9n-6n-3Fatty Acid Percentage of Seed Oil*Fatty AcidsComponent Fatty Acids in Major Asteraceae Oil-Seed Crops*et al.Iva annua Iva annuaHelianthus annuusCarthamus tinctorius
 Component Fatty Acids in Major Asteraceae Oil-Seed Crops
 This shows the compar-ison of the fatty acid composition of the oil from cypselae of 
Iva annua, 
Helian-thus annuus, and 
Carthamus tinctorius. The ratio of the n-6/n-3 fatty acids is accepted as 
n-3 FAs to form a preferred ratio.
106the cultivated (saline and fresh) and the wild (saline and fresh), although there is a down-regulated performance in the C 18:1 fatty acid levels for both cultivated in freshwater alternatives in contrast to the wild-collected samples. GC-FID with appropriate standards. It is possible that the Asch and Asch sample had unusually high 
levels of free fatty acids. If Asch and Asch had anomalous samples that were not representative in 
this characteristic, this would also show as lowered N content, perhaps pointing to some hints that could resolve the discrepancy our disparate values, both in lipids and in protein as well.High levels of free fatty acids are often not well tolerated in the physiology of seeds because of the deter-gent activities of these molecules, and their concomitant action in damaging membranes. Note, their simi-larity to sodium dodecyl sulfate (lauryl= C12). Sodium dodecyl sulfate, also called sodium laurilsulfate or 
sodium laryl sulfate is a surfactant (CH3(CH2)11SO4Na) used in various cleaning and hygiene formulations.for a number of health issues, often having to do with conditioning the heart in some way. A prevail
-because the human body cannot synthesize them, and valuable because they share the enzymes that 
this is that they function with their best effect when they are present in a nearly 50/50 ratio. We take 
107
synthesis of eicosanoids, a family including leukotrienes and prostaglandins. These act as cell mes
-
in wound healing and infection management, and too much of the n-3 family tends to support the im-balance (Dubois et al. 2007).The Principal Component Analysis shown in Figure 5-10 does not show any obvious consistent 
characteristics of the cypselae from the Wickliffe, Kentucky site (cult) show great dispersal within the 
site that would be consistent with great genetic diversity; well resolved on the X-axis See A in Figure 
5-9. When one is observing plants grown in the hypersaline conditions of the prairie salt marshes, 
one of the most obvious effects on 
I. annua, is the limited biomass of the plants; about half the height of the same species™ individuals from freshwater sites. The lack of strong evidence of NaCl-induced 

those grown in controlled saline environments.Data from the total lipid analysis reveal that cypselae weights are inversely correlated with per-cent lipid content. Because of their larger surface area to volume ratios, smaller seeds usually have 
comparatively high cell wall content which would tend to lower the lipid content. So, these small 108seeds with large oil percentages are interesting. Plotting the correlation between cypselae weight and 
lipid percentages yields the linear regression shown in Figure 5-10. This regression is striking, and 
apparently unprecedented. It implies that 97 percent of the variance of percent lipid is explained by of prediction of the lipid levels is within 3 percent. In soy, 
Glycine max, one of the most intensely researched oil seeds, no such correlation has been found (Maestri et al. 1998). The trend in larger 
10-70-60-50-40-30-20-1001020-4-202468Heiser INWickli˜e KY Site BLexington SCGranite City IL Site BWickli˜e Site AWickli˜e Site A Cult.P˚zer Large (single pl.)P˚zer SWLittle Salt Fork NE Cult.P˚zer SW Cult. Fert.P˚zer SW Cult.Lincoln SWSchoemaker SW CultivatedWild CollectedFresh waterSalt water1st Principal Component (96.87 percent of variability)2nd Principal Component (2.5 percent of variability)Principal Component Analysis of Fatty Acidsn=3n=3n=3n=3n=3n=6n=5n=8n=2n=3n=8
n=2
n=3
 Principal component analysis of the fatty acids
 
in FA composition differences among populations.
A109Figure 5-12
  Percent Nitrogen by Cypselae Weights
 

 Percent N
Figure 5-11 
 Percent Lipid by Cypselae
 Weights
 Sample weight vs. percent lipid showing the un
-
2Sample Weight mg (5 kernels)
110kernels, with lower oil levels, point to higher protein and/or starch levels. Comparing N content with seed weight (Figure 5.11) also indicates an inverse correlation with the exception of the two unusu
-ally low weight low N samples (omitted) pointing to starch levels being higher in larger seeds. 
Another line of investigation to test whether there is more to the saline success of I. annua than simply recent acclimatization is to test the performance of germinating seed collected from both saline and freshwater environments under various saline and freshwater conditions. It is possible that the populations of I. annuahave done so long enough to have made adaptation to coping with the ionic concentrations they face there. A test was performed to see if the endemic saline-inhabiting populations exhibited any special
-ized abilities during germination in differing concentrations.
I. annuathe Mississippi and northward sporadically. It has mostly been considered a coastal or riparian spe
-cies except for certain areas of Nebraska and Kansas where it has been noted as a regular inhabitant of the outer zones of salinity surrounding saline wetlands, and salt marshes. The salt tolerance of 
I. 
annua has been noted and described in some detail by Ungar and Hogan (1970). They describe the salt 
toleration as moderate and give the range of that tolerance as from 0.1 percent to 1.3 percent total salts 
(salinity). Although the authors use cypselae collected in the saline wetlands surrounding Lincoln, Ne
-braska for this study, they do not discuss whether these populations differ intrinsically from the popu
-lations that are found across the majority of non-saline conditions that comprise the natural range. To 
determine whether differences in seed salt tolerance between populations are due to genetic or environ
-mental effects, it will be necessary to compare the germination of seeds from plants representing saline 
111populations and fresh water populations that had been grown under controlled salinity conditions.and the greater germination success at the increased salinities implies that there is some degree of adaptation to the saline environment (Figure 5-8B, 5-8C). It is worth noting that Ungar and Hogan (1970) state that I. annua from saline wetlands inhabit soils averaging salinities of 0.5-0.7 percent total salts, while as reported here, germination was certainly not zero even at 1.8 percent salinity. 
There would be an expectation that while germination and survival can be accomplished at higher ability to compete with more accomplished halophytes that commonly inhabit the increased salinity areas of the habitable zone for I. annua in these wetlands. Ungar and Hogan say (Ungar and Hogan 1970, page 153), fiThe reduction in germination that takes place with increased salt concentration is not a permanent inhibition. . .This tolerance of I. annua to salinities up to 23% NaCl permits survival during dry periods, when the salinity hazard rises (Ungar 1968), and during periods of temporary ConclusionsThe use of I. annua seeds as food is one of the more enigmatic resources known. At the time of Eu
-ropean contact, it had either fallen out of use completely, or been retained by the smallest handful of 
American Indians. After Asch and Asch (1978) published their description of the economic potential 
of I. annua, it became apparent that this was a species that at one time had contributed substantially to the nutritional wellbeing of a large number of Native Americans east of the Mississippi. Certainly, 
112some of the reasons for this appreciation had to do with the oil and protein levels to be found in the present day cypselae as presented here. It seems reasonable to expect that these levels may be similar to the seed stock found in archaeological discovery. The fact that the protein content had been appar
-ently under-reported in the one analysis (Asch and Asch 1978) that preceded this investigation makes 
the estimated contribution of I. annua 
Why would cultures, as subject to the challenges of food abundance and climatic perturbations 
as those of ancient North America, give up this crop? In the book, 
Guns, Germs, and Steel Jared 
Diamond (1997, page 151) says a huge disadvantage of growing marshelder was its cross-sensi-tivity with  the hay-fever-causing ragweeds and that it was irritating to the skin. However, histori
-cally to attempt to validate the general perception that American Indians did not generally suffer 
hay-fever, Arthur F. Coca 
et al. (1922) found in interviews with physicians from American Indian 
schools, and serum tests with full-blood American Indian volunteers that in fact allergies were 
extremely rare in American Indians when compared with the those of European descent The sum
-

-tal study of the occurrence of serum disease in twenty-six volunteer full-blood American Indians 
indicates that the Indian race is much less susceptible to that condition than is the white race.fl 
Later, this comment appears in a letter to the New England Journal of Medicine in summary of 
consultations made with American Indian health workers, (Herxheimer 1964); fiI conclude that 
bronchial asthma was indeed almost unknown in American Indians before 1931 and that there is 
now an appreciable, but small number of typical cases in some tribes of Arizona and New Mexico 
113(Papagos, Hopis, Pimas, Zumis and Navajos) Πmuch less than in the white population.fl It seems in 
the case of a population of people that carefully nurtured a crop for more than 3 millennia, that they given it up for cause long ago. At the time when the Eastern Agricultural Complex crop species were in decline amongst the 
American Indian groups in Eastern North America, a different but productive suite of crops was tak
-ing hold. Several of the crops later grouped fiThe Three Sistersfl were beginning to be (Wagner and 
Carrington 2014) grown, including maize. However, recent discoveries (Sidell 2008) in archaeology 
show the comparative declines in the culture of I. annua and the rise of Z. mays, were played out as our understanding of how recently I. annua culture was maintained (Wagner and Carrington 2014) 
has extended into much more recent times than formerly appreciated.There seems to be an underlying assumption in the discipline of archaeology that the proven pres-Zea mays in the diets of Native Americans, as documented through the under
-standing of maize phytoliths (Asch Sidell 2008, Hart et al. 2007, Boyd and Surette 2010) also proves maize cultivation. My problem with the resulting desire to use this data to push back the dates of maize culture, as against cultivation is the disregard for the lack of supporting macrofossil evidence; cobs etc. Part of this point of view includes an implicit assumption that simply trading in maize, 
-sistent with our ideas (uncorroborated by evidence) of what primitive agricultural trade could have 
114Since the ultimate goal of growing crops could be presumed to prevent the starvation of the maxi
-mum number of the community population, then the high yielding maize crop, even though not 
competing with marshelder in the world of high nutrition, could have, by sheer calories, promised 
the increase in survival and reduced selection pressure from hunger, that is an underlying purpose 
for the agricultural endeavor.
Finally, agriculture is wrapped into the cultural realm of fashion as are most human endeavors. 
After the comparative prosperity of maize or three sisters agriculture had been realized, there 
may have been social pressures to make the move to a more ‚modern™ selection of crop plants. 
Given the newly presented attributes of Iva annua, perhaps some future attempt might be made 
to more closely investigate whether this species should again be looked at in the world of agricul-ture. As a candidate for modern domestication it does present certain advantages and issues. On 
the ‚plus™ side the huge potential yields of protein, for a food seed are impressive. Even though 

-6 and low in -3 fatty 
acids, lipids still represent a value dietary and agricultural commodity. The potential value of 
raising such a crop on marginally salt contaminated land, would be welcomed in many parts of 

of Iva annua strongly cross-sensitizes with that of its close relatives in the genus 
the 

Iva annua is a troublesome 
weed over a considerable part of its natural range in the southern United States. One could easily 
imagine this plant, if introduced into other regions, escaping into the surrounding countryside 
with economically challenging results. Another close relative 
Cyclachena xanthifolia (formerly 
115Iva xanthifolia) has escaped in Europe into agricultural lands with undesirable effects (Weber 
and Gut 2005). More research is needed to complete these assessments of its potential value to our 
species.116REFERENCES117REFERENCESASCH, D. L., A
SCH, NANCY B. (1978). fiThe economic potential of 
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fl 
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BRODY, T. (1999). Vitamins. 
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EVERSON, G. M. (1950). fiThe Estimation of Oil Percentage in   

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