Phragmites use by Native North Americans

Aquatic Botany 69 (2001) 341–357

Phragmites use by Native North Americans Erik Kiviat∗ , Elizabeth Hamilton Hudsonia Ltd., Bard College, P.O. Box 5000, Annandale, NY 12504-5000, USA

Abstract Phragmites australis (Cav.) Trin. ex Steud. occurred in North America in pre-Columbian time but has spread greatly and become invasive in the north-central and northeastern US during the 1900s. At least 46 Native North American tribes were reported to use Phragmites historically, and we compiled statistical data from the literature on 24 of these tribes. There were approximately 75 different uses of Phragmites. Each tribe had 2–25 (mean = 7.0) uses for Phragmites, of which the most frequent were: arrowshaft (17 tribes), cigarette (13), flute (12), whistle (7), pipestem (7), and matting (6). Number of uses per tribe was highest in the SouthWestern quadrant of North America, and was negatively correlated with latitude but not correlated with longitude. The apparent center of Phragmites availability and use in the SouthWestern United States is consistent with occurrence of Phragmites in pre-Columbian sediments and archaeological deposits, and at least two specialized Phragmites-using animals. The ethnobotanical importance of Phragmites should be considered in management of this plant. © 2001 Elsevier Science B.V. All rights reserved. Keywords: Phragmites australis; Poaceae; Invasive species; Ethnobotany; Native Americans; North America

1. Introduction Phragmites australis (Cav.) Trin. ex Steud. (hereinafter “Phragmites”) is one of the most widely distributed vascular plants in the world (Clevering and Lissner, 1999). The only species of Phragmites in North America, P. australis is widespread and locally abundant in southern Canada, much of the United States, and regions of Mexico. Phragmites abundance has increased greatly in many areas of the north-central and northeastern US during the 1900s (Sipple, 1972; Marks et al., 1994; Stalter and Baden, 1994; Buck, 1995; Rice and Stevenson, 1996; Winogrond and Kiviat, 1997; Pellegrin and Hauber, 1999; Chambers et al., 1999). This invasion may be due to recently introduced subspecific forms, or to human-caused changes in the environment (Chambers et al., 1999). ∗ Corresponding author. Tel.: +1-845-758-7273; fax: +1-845-758-7033. E-mail address: [email protected] (E. Kiviat).

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Phragmites colonizes moist, saturated, or flooded soils that have been altered or disturbed. These habitats include filled and drained wetlands, deposits of dredged material, impounded tidal salt marshes, mine pits, farm ponds, ornamental and urban ponds, sewage ponds, constructed wetlands, highway verges and ditches, drainage or irrigation ditches, wetlands polluted, silted, or damaged by road or pipeline construction, post-agricultural wet meadows, abandoned rice fields, abandoned beaver (Castor canadensis) ponds, river banks, and areas of brackish or freshwater tidal marshes with elevations near Mean High Water (Martin et al., 1957; Wheat, 1967; Rea, 1983; Kiviat, 1987 and personal observations; Sinicrope et al., 1990; Marks et al., 1994; Stalter and Baden, 1994; Buck, 1995; Havens et al., 1997; Mathis and Middleton, 1999). Phragmites has also been planted as a garden ornamental (Duke, 1992) and in systems constructed for waste treatment (Hammer, 1992). Natural habitats of Phragmites apparently included the landward margins of tidal salt marshes (Orson et al., 1987; Orson, 1999), brackish marshes of river deltas (Castetter and Bell, 1951), alkaline or saline inland wetlands (McVaugh, 1958; Stewart and Kantrud, 1969) and river banks (Castetter and Bell, 1951), oases (Rodr´ıguez-Estrella et al., 1999), seeps (Rea, 1983), springs, and marshes in arid lands. Botanical manuals have generally treated P. australis as native in North America (Kearney and Peebles, 1951; Gleason and Cronquist, 1991; Munz and Keck, 1963; Seymour, 1969; Long and Lakela, 1971). A.E. Schuyler (Academy of Natural Sciences, Philadelphia, PA, pers. comm.) suggested that P. australis was brought to the northeastern US as an economic plant by early Dutch settlers, and Bernd Blossey (Cornell University, Ithaca, NY, pers. comm.) suggested that P. australis was brought to North America by pre-Columbian people crossing the Bering Sea from Asia. To explain the recent invasiveness of Phragmites in the northeastern states, it has been hypothesized that there are non-invasive native genotypes and invasive introduced genotypes of P. australis (Tucker, 1990; Marks et al., 1994). Available genetic data are insufficient to clearly distinguish invasive forms from non-invasive forms, however, and changes in wetland environments could also explain the recent invasiveness of this species (Chambers et al., 1999). In North America, Phragmites has leafy culms 1–2.5 cm in diameter and 1.5–5 m tall at maturity (Felger and Moser, 1985). The culms arise from extensive rhizomes 1.5–2.5 cm in diameter. The culms, usually erect, have long hollow internodes and diaphragms at the nodes, and are harsh, tough, and resistant to decay. Phragmites may attain a peak aboveground biomass of 1500–4000 g m−2 dry weight (Whigham et al., 1978; Meyerson et al., 2000). These characteristics, combined with local abundance, make Phragmites useful to humans. We undertook this study to assess the overall importance of Phragmites to Native North Americans, examine relationships between use of Phragmites and its geographic distribution, and determine if ethnobotanical data would clarify the past distribution of Phragmites in North America.

2. Methods We examined literature and selected unpublished data on the North American Indians for uses of Phragmites. Ethnographic identification of Phragmites is confused by a variety of common names including reed, common reed, reed grass, common reed grass,

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cane, common cane, wild cane, arrow cane, cane grass, carrizo, carrizo grass, carrizal reed, roseau, and bent grass. Some of these names have also been applied to Scirpus, Typha, Acorus, Arundinaria, and other taxa. Furthermore, synonyms for P. australis found in the ethnobotanical or ethnographic literature include P. communis, P. communis var. berlandieri, P. phragmites, P. vulgaris, and Arundo phragmites (see additional synomyms in Hitchcock, 1971). Therefore, our ethnobotanical data set is based on literature references to “Phragmites” (or “Arundo phragmites”) with the following exceptions. Pawnee used hollow “reeds” (Murie, 1981) which we interpreted as Phragmites. Many western tribes used “cane” and its identity was clear from intermittent use of the name “Phragmites” in the literature. Arundinaria, also called cane, and also hollow, does not occur in the regions in question. For the purposes of this paper, we assume that all Phragmites reported in North America in ethnobotanical, archaeological, and palaeoecological sources was P. australis because we are aware of no other species of Phragmites ever having been identified here (Clevering and Lissner, 1999). Tribes included in our statistical sample were all reported to use Phragmites historically (i.e. in post-Columbian time). We strove for consistency of interpretation of uses across tribes despite incomplete information. For example, as many as eight unspecified types of baskets made by the Pima included Phragmites material (Tanner, 1990) but we counted these as a single “use” because similar detail was not available for other tribes. Yet we counted several types of musical instruments made of Phragmites because these were clearly distinguished in the literature, e.g. whistle, end-blown flute, and flageolet, all made by Apache (Gifford, 1940). Our use of the term “tribe” indicates an entity identifiable by culture for the purposes of our study; we do not imply a particular sociopolitical structure. We mapped a centroid of the area occupied by each tribe to the nearest 0.5◦ of latitude and longitude, and assigned the tribes to compass quadrants separated by 36◦ north latitude and 100◦ west longitude. Our term “Southwest” includes the southwestern US and northwestern Mexico. For statistical analysis, we used the Kruskal–Wallis one-way analysis of variance by ranks (H) and the Spearman rank correlation (rho) because data violated assumptions of normality, linearity, and homogeneity of variances. Because our data set contained only one southeastern tribe, we removed the southeastern quadrant from an analysis of differences among quadrants. Statistics were computed with Statistica version 5.9 (StatSoft., 1995). Spatially close tribes may be non-independent sampling units due to propagation of culture through space (Burton and White, 1987; Mace and Pagel, 1994). Therefore, we performed separate statistical analyses on a complete data set of 24 tribes, and a reduced data set of 19 tribes after randomly removing Apache, Havasupai, Maricopa, Northern Paiute, and Papago from clusters of southwestern tribes. If near neighbors belonged to different language branches (Champagne, 1994) or had substantially different economies (e.g. foraging versus cultivation), however, we retained all tribes in the reduced set.

3. Results In our statistical sample, Phragmites use is widely distributed in North America, but with a concentration of tribes and a greater diversity of uses in the Southwest (Fig. 1, Table 1).

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Fig. 1. Locations of 24 Phragmites-using Native North American tribes in the statistical sample (refer to Table 1 for names of tribes) (map prepared by K. Cafasso and G. Stevens).

Our data include approximately 75 types of uses (Table 2 ), the number somewhat dependent on judgment used in classification within and across tribes. Number of uses per tribe is 2–25 (mean = 7.0, median = 4.5) (Table 1). All plant parts were used, but the culm was most important by far. The most frequent uses of Phragmites were arrowshaft (17 tribes), cigarette (13), flute (12), whistle (7), pipestem (7), matting (6), flageolet (5), and knife (5). Phragmites was used for arrowshafts in three compass quadrants but mainly in non-forested environments (Chippewa was an exception) and mainly in the Southwest (Fig. 1). Smoking uses occurred in two quadrants and were mainly southwestern. “Chemical” uses (medicine, seed soak) occurred in all four quadrants. Matting, basketry, and cordage uses occurred in three quadrants but were all western except Chippewa. Musical instruments occurred in three quadrants but were southwestern except Iroquois and Pomo. Game pieces and knife were exclusively southwestern (one quadrant). Sugar occurred only in the Southwest (one quadrant), but outside our statistical data set Phragmites was widely used for sugar. Sugar was derived from Phragmites sap as well as from the honeydew of aphids, probably Hyalopterus pruni Geoffr., on Phragmites (Heizer, 1945; Jones, 1945; Wheat, 1967). In the complete data set (n = 24), number of uses differs among the three compass quadrants (Fig. 1, Table 1) (Kruskal–Wallis H = 13.9, P = 0.0010). Number of uses is higher in the Southwest (median = 9.5, range = 3–25) than in the Northeast (median = 2, range = 2–2) or Northwest (median = 4, range = 2–4). Number of uses is negatively correlated with latitude (rho = −0.453, P = 0.0260) and is not correlated with longitude (rho = 0.294, P = 0.163). Probabilities for rho and H are virtually the same for the reduced data set.

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Table 1 Phragmites-using Native North American tribes in the statistical sample (n = 24) (see Fig. 1 for locations) Tribe

Map no. Longitude Latitude Uses References

Apache Blackfoot Cahuilla

1 2 3

106.0 112.0 117.5

33.5 49.0 35.5

14 3 6

Chipewyan Chippewa Chumash

4 5 6

102.0 94.0 120.0

61.0 47.0 34.5

2 2 11

Cocopa Gosiute Havasupai Hopi

7 8 9 10

115.0 114.0 112.0 110.5

32.0 39.5 36.0 36.0

3 4 3 11

Hualapai Iroquois Kawaiisu Klamath Maricopa Navajo

11 12 13 14 15 16

113.0 78.0 118.5 121.5 112.5 108.0

35.0 43.5 35.5 42.5 33.5 36.0

5 2 13 4 7 8

Northern Paiute 17

118.5

41.5

15

Papago

18

112.0

32.0

7

Pawnee Pima

19 20

99.5 112.0

41.0 32.0

2 12

Pomo Seminole Seri Thompson

21 22 23 24

123.0 81.0 112.0 121.0

39.0 26.5 29.0 50.0

2 2 25 4

Reagan, 1929; Gifford, 1940; Duke, 1992 Johnston, 1970, 1987 Barrows, 1967; Bean, 1972; Bean and Saubel, 1972; Heizer and Elsasser, 1980 Marles, 1984 Lyford, 1943; Petersen, 1963; Densmore, 1974, 1979 Timbrook, 1990; Jan Timbrook, Santa Barbara Museum of Natural History, pers. comm. Drucker, 1941; Castetter and Bell, 1951; Kelly, 1977 Chamberlin, 1911; Stewart, 1942; Steward, 1943 Weber and Seaman, 1985 Gifford, 1940; Whiting, 1966; Dunmire and Tierney, 1977 Drucker, 1941; Watahomigie et al., 1982 Waugh, 1916; Chafe, 1967; Herrick, 1995 Driver, 1937; Zigmond, 1981 Coville, 1897; Barrett, 1910; Spier, 1930 Drucker, 1941; Castetter and Bell, 1951 Reagan, 1929; Gifford, 1940; Stewart, 1942; Elmore, 1943; Dunmire and Tierney, 1977 Stewart, 1941; Train et al., 1957; Wheat, 1967; Fowler, 1989, 1990 Castetter and Underhill, 1935; Gifford, 1940; Drucker, 1941; Castetter and Bell, 1942 Murie, 1981 Drucker, 1941; Castetter and Bell, 1942; Cain, 1962; Russell, 1975; Curtin, 1984; Tanner, 1990; Rea, 1997 Barrett, 1952 Sturtevant, 1955 Dawson, 1944; Felger and Moser, 1985 Steedman, 1928; Turner et al., 1990

4. Discussion 4.1. Geography of Phragmites use Among Native North Americans, Phragmites australis was used more than most herbaceous plants, but the number of uses and the number of user tribes were substantially less than for the most highly used herbaceous plants and the more highly used woody plants (Moerman, 1998). Phragmites was important to many tribes in the Southwest and probably was a critical resource for arrowshafts, sugar, smoking equipment, and certain other uses. In addition to the 24 tribes in our statistical data set, Phragmites use has been reported for at least 22 other tribes: Achomawi (Merrill, 1923; Stewart, 1941), Akwa’ala (Drucker, 1941), Cochimi (Heizer, 1945; Heizer and Massey, 1953), Costanoan (Heizer, 1945), Diegueño (Drucker, 1941; Shipek, 1993), Lillooet (Turnbaugh and Turnbaugh, 1986), Mohave (Drucker, 1941), Northern Shoshoni (Steward, 1943), Panamint (Coville,

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Table 2 Uses of P. australis reported for the Native North American tribes of the statistical sample (n = 24) Use

Tribes

Food Seed Sugar Young shoot

Klamath Chumash, Gosiute, Kawaiisu, N. Paiute Chipewyan

Fishing and hunting equipment Arrowshaft Call (plunged in water to attract fish) Call (for rabbit) Mats, baskets, cordage Basketry Cordage, unspecified Cordage carrying net Cordage baby hammock Mat for drying food Mat (sleeping) Mat (table) Mat (unspecified) Musical instruments Arrow to play musical bow Clapper Flageolet Flute Mouth bow Rattle Whistle

All tribes except Chipewyan, Iroquois, Pawnee, Pima, Pomo, Seminole, Thompson Seri Seri Klamath, Pima, Thompson N. Paiute Cahuilla Cahuilla Chippewa, Havasupai, Thompson Pima Pima Maricopa, Seri Kawaiisu Kawaiisu Apache, Hualapai, Maricopa, Papago, Pima Apache, Cahuilla, Cocopa, Hopi, Hualapai, Iroquois, Maricopa, Navajo, N. Paiute, Papago, Pima, Seri Seri Pomo, Seri Apache, Chumash, Hopi, Navajo, N. Paiute, Pomo, Seri

Shelter construction Door covering Frame material “Roofing” Thatch

Pima Seri Hopi Cahuilla, Chumash

Clothing and personal adornment Apron Basketry hat Beads for fringe of dress Ear ornament Necklace

Chumash Klamath Thompson Kawaiisu, N. Paiute N. Paiute, Thompson

Games “Bones” Counting stick for games Dice Ring and pin game Tube for flipping shell into Tubes to hide object in (guessing game)

Kawaiisu, N. Paiute Chumash, Seri Gosiute, Kawaiisu, Maricopa, Seri Kawaiisu Seri Maricopa, Papago, Pima, Seri

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Table 2 (Continued) Use

Tribes

Ceremonial items Cloud blower Flute used by Shaman Frame for Mountain Chant Medicine bundle (Big Black Meteoric Star) Medicine bundle (Wonderful Leggings) Object associated with bow and arrow Prayer stick Twirling wand Walking stick used by Shaman

Pima Seri Navajo Pawnee Pawnee Hopi Hopi, Navajo, Pima Pima Hualapai, Pima

Smoking equipment Cigarette (tubular pipe)

Pipestem Container for tobacco storage

Apache, Chipewyan, Chumash, Cocopa, Hopi, Hualapai, Kawaiisu, Maricopa, Navajo, N. Paiute, Papago, Pima, Seri Apache, Blackfoot, Gosiute, Havasupai, Hopi, Navajo, N. Paiute Apache, Chumash

Medicine “Root”, for digestive problems Sugar, for pneumonia Unspecified part, for boils or carbuncles Whole plant, as emetic

Apache N. Paiute Seminole Apache, Blackfoot, Navajo

Medical equipment Medicine blowing tube Splint for fractures Tube for magical powder of Shaman Tube for suction by Shaman

Seminole Cahuilla, Chumash Seri Apache, Papago

Other Canoe Container for pollen or pigment Drinking straw Fire drill shaft Fire starter (charcoal inside culm) Fruit gathering pole Knife (various uses) Loom of perforated culms Mnemonic devices Paintbrush handle Soaking seed maize Toy harpoon Toy (pea shooter) Toy (popgun) Toy gun for shooting slivers Toy (squirt gun) Tweezers for beard Vent for cooking coyote Walking stick for elderly Weaving rods

Seri Seri Apache, Seri Kawaiisu, N. Paiute N. Paiute Seri Chumash, Hopi, Kawaiisu, N. Paiute, Seri Hopi Apache Chumash Iroquois Seri Apache, Seri Papago Seri Seri Kawaiisu, N. Paiute Kawaiisu Apache Hopi

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1892), Pueblo (San Ildefonso, Santa Ana, Walpi, Zuñi) (Gifford, 1940), Quinault (Turnbaugh and Turnbaugh, 1986), Salish (Laforet, 1990), Shuswap (Turnbaugh and Turnbaugh, 1986), Southern Paiute (Stewart, 1942; Heizer, 1945), Tewa (Robbins et al., 1916), Ute (Gifford, 1940; Dunmire and Tierney, 1977), Washo (Stewart, 1941), Yaqui (Drucker, 1941), Yavapai (Drucker, 1941), Yokuts (Driver, 1937; Heizer, 1945), Yuma (Castetter and Bell, 1951), and at least one unspecified tribe in the southern Appalachian Mountains (Core, 1967). These additional tribes were in northwestern Mexico, southwestern, northwestern, and southeastern US, and southwestern Canada. The compilation of North American Indian ethnobotany by Moerman (1998) reported Phragmites use by 22 of 290 tribes or 7.6% of the tribes for which data were compiled. This percentage is conservative because lists of plants used by a tribe are frequently incomplete. Historical loss of ethnobotanical knowledge, and the limits to our survey, suggest the full number of Phragmites-using tribes may have been much greater than 46. P. australis had widespread, often intensive, uses in Europe (Rodewald-Rudescu, 1974) and was at least a locally important economic plant in Africa (Rodin, 1985, p. 83), South America (Heiser, 1979), and Australia (Zola and Gott, 1992). Duke (1992) listed a variety of uses in folk medicine worldwide. A closely-related species, Phragmites karka, was the basis of the material culture of Madan in Iraq and Iran (e.g. Salim, 1962; Thesiger, 1964), the most Phragmites-dependent culture in the world. Non-industrial use of Phragmites in the Old World bears comparison to North America. Use as human food in Europe was uncommon, and we are not aware of any report of sugar collection from Phragmites outside North America. P. australis has been used for arrowshafts by San in Namibia (Tim Abbott, Nature Conservancy, Sheffield, MA, pers. comm., 1999), and for spearshafts by Koorie in Australia (Zola and Gott, 1992); P. karka (Retz.) Trin. ex Steudel has been used for spearshafts by Madan (Thesiger, 1964). P. australis has been used widely for thatching in Europe; this use is rare in our statistical sample but was probably widespread in the Southwest. Boat construction from Phragmites apparently had a very local distribution around the world (e.g. Heizer and Massey, 1953; Thesiger, 1964). The importance of a plant species to a society may be assessed by the development of culture associated with collection and processing, as well as by the diversity of uses. The variety of methods by which sugar was produced from Phragmites attests to its importance in regions lacking sugar-producing trees (especially Acer saccharum). Phragmites sugar was made by drying and grinding the whole shoots then roasting the resulting “flour” (Heizer, 1945); allowing harvested shoots to wilt or dry, then beating them and collecting the small, sweet particles so detached (Heizer, 1945; Jones, 1945; Wheat, 1967, p. 116; Zigmond, 1981, p. 49); drying and eating the leaves (Steward, 1943, p. 302); and scraping sweet material off the leaves (Palmer, 1878; Steward, 1943, p. 302). Other indicators of the importance of Phragmites include effort expended to obtain Phragmites and specialized equipment for collecting or processing Phragmites. A Seri group had to travel for several days to collect Phragmites for various uses (Felger and Moser, 1985). Zuñi imported Phragmites for flutes (Gifford, 1940). Northern Paiute made a special basket for gathering Phragmites sugar (Fowler, 1990). Some tribes had a special tool and procedure for straightening Phragmites arrowshafts (Heizer and Elsasser, 1980, p. 141). Yavapai made a “wrench” specifically for tightening the junction of the Phragmites mainshaft and wood foreshaft of an arrow (Gifford, 1936, p. 36). Diegueño burned stands of Phragmites to improve its quality for

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arrowshafts (Shipek, 1993). Lastly, ceremonial and religious uses of Phragmites (Table 2) indicate its cultural importance to certain tribes. 4.2. Alternate plants The technological importance of Phragmites in non-industrial settings is related to the characteristics of the plant, especially, the culms, which are strong, hollow, lightweight, buoyant, rapidly growing, and slow-decomposing. These characteristics, and hence many of its uses, are unusual for an herbaceous plant. Some uses of Phragmites (matting, basketry, young shoots as food, boat construction) are similar to uses of Scirpus spp. and Typha spp., whereas other uses (arrowshaft, fire drill shaft, splint) are related to special qualities of Phragmites rarely found in other robust marsh graminoids. Phragmites use was widespread in North America but evidently concentrated in the Southwest. We are puzzled by the absence or scarcity of ethnobotanical data from coastal Louisiana where Phragmites is prominent (Lynch et al., 1947; Pellegrin and Hauber, 1999), and coastal New England where Phragmites fossils have been recovered from a number of tidal marshes (Orson, 1999). We think the pattern shown in Figs. 1 and 2 is due to a combination of actual intensity of use and adequacy of ethnobotanical observation. That there is evidence of few Phragmites-using tribes, and few uses of Phragmites per tribe, in eastern North America could be due to: (1) less Phragmites present until recently; (2) eastern tribes did not adopt uses of Phragmites, or uses were not documented, before loss of Native American culture; (3) eastern ethnobotanists did not ask Native Americans about Phragmites because it was considered a recently introduced species (Roger Moeller, Archaeological Services, Connecticut, pers. comm., 1999); or (4) there were better plants available for particular uses. In the case of plants used for basketry by Native Americans in California, the geographic distribution of certain species (including Phragmites) greatly exceeded the distribution of use, whereas the distribution of other species but little exceeded the distribution of use (Merrill, 1923). Our data indicate that uses of Phragmites varied from widespread (e.g. arrowshaft) to local (e.g. basketry). In the Southwest, however, Phragmites was an important economic plant for many tribes. Outside the arid Southwest, there may have been greater variety and quantity of alternate materials for human use. Pomo, in a partially forested region of northern California, had only two uses of Phragmites (Tables 1 and 2). In the eastern states, many woody plants were used for arrowshafts, and substitutes such as Arundinaria gigantea Michx. (e.g. Kniffen et al., 1987) may have lessened the importance of Phragmites. Nonetheless, a shrub, Pluchea sericea (Nutt.) Coville was used alternately with Phragmites for arrowshafts by Pima and other southwestern tribes (Drucker, 1941, p. 118; Heizer and Elsasser, 1980, p. 141). Apache, Gosiute, and Navajo used the stout, pithy twigs of Sambucus, a woody plant, interchangeably with hollow Phragmites internodes for pipestems, and Southern Paiute and Ute used Sambucus interchangeably with Phragmites for cigarettes (Gifford, 1940, p. 57; Stewart, 1942, p. 293). Apache used Sambucus or Phragmites for drinking straws (Gifford, 1940, p. 65). Kawaiisu and Panamint used wood or split Phragmites alternately for tweezers (Driver, 1937, p. 73). Honeydew for sugar was collected from Salix gooddingii Ball, Populus angustifolia James, Pluchea sp., and Scirpus lacustris L. (Scirpus acutus Muhl. or S. tabernaemontani K.C. Gmel.), but Phragmites was evidently the most widespread sugar

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Fig. 2. Scatterplots of number of Phragmites uses per tribe against latitude and longitude for 24 tribes in the statistical sample.

plant in the Southwest (Heizer, 1945; Castetter and Bell, 1951, p. 218). Arundo donax L., native in southern Europe and introduced in the southern and western United States and northwestern Mexico, has been used interchangeably with Phragmites for certain purposes on both continents, including by Cahuilla, Chumash, Hopi, and Seri in North America (Whiting, 1966, p. 64; Bean and Saubel, 1972, p. 102; Felger and Moser, 1985; Timbrook, 1990, p. 246). 4.3. Human use and palaeoecology Utility of a plant to humans is related to: (1) abundance and distribution of the plant; (2) length of time the plant and a human group have been in contact; (3) invention or transmission of traditional ecological knowledge of the taxon; (4) ease of managing, acquiring, and

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processing the plant; (5) physical and chemical qualities of the plant (e.g. pharmaceutical or toxicological properties, fiber characteristics, nutritional composition); and (6) availability and quality of alternate taxa. We believe the relative variety and intensity of uses of Phragmites by a human group in North America indicate the length of time the people and the plant were in contact. Variety of uses has been considered an indicator of the length of history of human–plant interaction, e.g. coconut, Cocos nucifera L. (Zizumbo-Villarreal and Quero, 1998). We hypothesize that Phragmites was more abundant and widespread in southwestern North America during wetter Pleistocene or Holocene climatic intervals, especially after Pleistocene mega-herbivores became extinct, and before cattle, horses, and sheep were widely introduced. Intensive use of Phragmites by Native Americans may have developed during the period approximately 1450–600 years B.P. which is represented by many Phragmites artifacts in archaeological sites (e.g. Adams, 1990; unpublished catalog data provided to Kristin Saltonstall by Mike Jacobs, Arizona State Museum, Tucson, AZ, and Gian Mercurio, Mesa Verde National Park, CO). Native Americans might have dispersed or planted Phragmites, as suggested by Dunmire and Tierney (1977) and Gian Mercurio (Mesa Verde National Park, CO, pers. comm., 2000). Native North Americans effected intentional or unintentional dispersal or propagation of many other useful, non-domesticated plants (e.g. Yarnell, 1964; Blackburn and Anderson, 1993). Following European settlement of North America and onset of a drier climate, a major decline and fragmentation of Phragmites populations may have been caused by introduced livestock (e.g. Twisselmann, 1967, p. 180; Rea, 1997, pp. 102–103; Rodr´ıguez-Estrella et al., 1999), intensive agriculture, removal of groundwater (Rea, 1983; Curtin, 1984, p. 75), hydrological disruption of rivers (Rea, 1997), increased salinity intrusion in tidal wetlands (Castetter and Bell, 1951), and destruction of wetlands. We think the plant was sufficiently important to Indians, however, that its use would have persisted. In the 1900s, Phragmites spread extensively and became invasive in north-central and northeastern North America. This population explosion was probably facilitated by greatly diminishing numbers of free-ranging livestock, reduced competition from other robust wetland herbs and woody plants in disturbed or artificial habitats, reduced salinity in impounded coastal salt marshes, and pollution of freshwater marshes by de-icing salts, fertilizers, sewage, and sediments. In situ evolution of greater invasiveness, or recent introduction of one or more invasive genotypes from another continent (Marks et al., 1994; Pellegrin and Hauber, 1999; Chambers et al., 1999), may have played a role. Phragmites appears most abundant in habitats with moderate conductivity, including brackish estuarine marshes, fens, wetlands and riparian habitats of arid lands, and polluted habitats (E. Kiviat, personal observations). Arid southwestern regions are characterized by soils (Aridisols) that frequently have accumulations of lime, gypsum, or salt, and high pH (Miller and Donahue, 1995). In early historic times, these arid regions may have had more Phragmites than many other regions of the continent. The apparent concentration of tribes using Phragmites in the Southwest, and the higher numbers of uses per tribe there, are consistent with this hypothesized distribution of Phragmites. The Southwest also has widespread occurrence of Phragmites in archaeological sites, both pre-Columbian (e.g. Kaplan, 1963; Aikens, 1970; Bohrer, 1973; Adams, 1990; Dello-Russo, 1999) and historic (e.g. Seymour, 1998). Phragmites of pre-Columbian age and possibly as old as 7000 years has also been

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found in sediments of the Sacramento — San Joaquin Delta of central California (Atwater, 1980). The oldest known North American Phragmites remains were recovered from coprolites of the extinct Shasta ground sloth, Nothrotheriops shastense Sinclair, in the Southwest (Arizona), dated from greater than 40,000 to ca. 11,000 years B.P. (Hansen, 1978). Further support for a pre-Columbian concentration of Phragmites in the Southwest or West is provided by two animals: a monophagous, Phragmites-feeding butterfly, the Yuma skipper, Ochlodes yuma (Edwards), endemic to the western United States (Scott et al., 1977), and a wetland bird, Belding’s yellowthroat, Geothlypis beldingi, closely associated with Phragmites in desert oases of Baja California, Mexico (Rodr´ıguez-Estrella et al., 1999). The conjunction of Phragmites fossils, remains and artifacts from archaeological sites, historic Indian uses of Phragmites, and Phragmites-associated animals in the Southwest suggests that there should be similar co-occurrence on the northeastern coast from at least Massachusetts to New Jersey. Pre-Columbian Phragmites fossils have been well-documented from Massachusetts to New York City (Orson et al., 1987; La Porta et al., 1999; Orson, 1999; Philip C. La Porta, La Porta and Associates, Warwick, NY, pers. comm., 2000). A carbonized, modified, Phragmites rhizome ca. 325 years old was identified by David Perry from a Native American archaeological site at the Mashantucket Pequot Reservation on the eastern Connecticut coast (Kevin McBride, University of Connecticut, Storrs, pers. comm., 1999). David Perry (Stanfordville, NY, pers. comm., 2000) also identified four Phragmites fragments from an archaeological site of similar age at Gay Head, MA. Coastal populations of a butterfly, the broad-winged skipper, Poanes viator zizaniae Shapiro, feed on Phragmites and Zizania aquatica in the northeastern states (Opler and Krizek, 1984), although it is unclear whether the relationship of the skipper to Phragmites is of ancient or recent origin. Occurrence of Phragmites fossils, Phragmites materials in early historic archaeological sites, and the existence of a stenophagous, Phragmites-feeding insect all suggest the antiquity of Phragmites on the northeastern coast. Phragmites was evidently common and characteristic here before the 1900s although apparently not occurring in extensive stands (see Orson, 1999). We predict that additional historic and prehistoric Indian use of Phragmites will be discovered in this coastal region. Ethnobotanical data are, by nature, of heterogeneous quality. It is difficult to know how complete the recording of uses of Phragmites has been for each tribe. In North America, collection of ethnobotanical data has been most thorough in the Southwest. This could have exaggerated the number of uses per tribe (relative to non-southwestern tribes) thus producing part of the pattern in our data which show more uses southward and westward. Further compilation of data collected from living cultures, archaeological sites, and artifacts in museums is needed to understand spatial and temporal patterns of Phragmites distribution and Phragmites use by Native Americans. Our synthesis relates the antiquity of occurrence of Phragmites in North America, its ethnobotanical importance to Native North Americans, and its association with at least two species of specialized animals. 4.4. Implications for management policy Plant distributions are continually in flux, due to natural factors and human activities (Yarnell, 1964; Blackburn and Anderson, 1993). Our analysis attempts to relate human use of Phragmites to the biogeography and ecology of Phragmites in North America. The

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importance of Phragmites to Native Americans raises ethical issues about efforts to manage Phragmites on a large scale or on lands used by Native Americans, for example, a proposed biological control program (Schwärzlander and Häfliger, 1999). Native basketweavers are intensely concerned about their ability to manage and collect traditional plant materials (Ortiz, 1993). Cultural revival in many tribes could increase the importance of Phragmites as an economic plant. Knowledge of ethnobotany is also relevant to a search for additional beneficial uses of Phragmites in industrial society. Finally, there are many efforts in ecological restoration of wetlands and riparian habitats, and increased knowledge of the prehistoric and historic geography of Phragmites can provide more accurate “targets” for restoration design incorporating locally characteristic plants and their human and wildlife users.

Acknowledgements We thank, for literature, information, ideas, and assistance, Tim Abbott, Karen Adams, Kat Anderson, Sumru Aricanli, Brian Atwater, Bernd Blossey, Mary Burns, Kristen Cafasso, Josh Collins, Robert Dello-Russo, Joe Diamond, Leslie Driscoll, Christian Feest, Stewart Fefer, Antonia Giardina, Ives Goddard, Katherine Gould-Martin, Peter M. Groffman, Deborah G. Harding, Mike Jacobs, Herbert Kraft, John Kraft, Philip C. La Porta, Tonya Largy, Chris Lindner, Jason Mancini, Cecilia Maple, Kevin McBride, Dareth McKenna, Lucinda McWeeney, Gian Mercurio, Laura Ahearn Meyerson, Kate Mini, Roger Moeller, Richard Orson, David Perry, Kristin Saltonstall, Ken Scarlatelli, A.E. Schuyler, Gregory R. Seymour, Nicholas N. Smith, Gretchen Stevens, William Sturtevant, Jan Timbrook, Amy Toth, Tom Vennum, the Hudson River National Estuarine Research Reserve, and the Bard College Library. Karen Adams, Hans Brix, Randy Chambers, Tonya Largy, Daniel Moerman, Jane Roberts, and Kristin Saltonstall provided helpful comments on a draft. The Wildlife Conservation Society funded concurrent studies of Phragmites. This is Bard College Field Station — Hudsonia Contribution 76. References Adams, K.R., 1990. Prehistoric reedgrass (Phragmites) “cigarettes” with tobacco (Nicotiana) contents: a case study from Red Bow cliff dwelling. Arizona J. Ethnobiol. 10, 123–139. Aikens, C.M., 1970. Hogup Cave. Univ. Utah Anthropol. Pap. 93. Atwater, B.F., 1980. Attempts to correlate late Quaternary climatic records between San Francisco Bay, the Sacramento — San Joaquin Delta, and the Mokelumne River, California. Ph.D. Thesis. University of Delaware, Newark. Barrett, S.A., 1910. The material culture of the Klamath Lake and Modoc Indians of northeastern California and southern Oregon. Univ. California Pub. Am. Archaeol. Ethnol. 5, 239–294. Barrett, S.A., 1952. Material aspects of Pomo culture. Bulletin of the Public Museum of the City of Milwaukee 20, Parts I–II. Barrows, D.P., 1967. The Ethno-botany of the Coahuilla Indians of Southern California. Malki Museum Press, Banning, CA (reprint of 1900 edition). Bean, L.J., 1972. Mukat’s People. The Cahuilla Indians of Southern California. University of California Press, Berkeley. Bean, L.J., Saubel, K.S., 1972. Temalpakh. Cahuilla Indian Knowledge and Usage of Plants. Malki Museum Press, Banning, CA.

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