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Typha latifolia

• INTRODUCTORY
• DISTRIBUTION AND OCCURRENCE
• BOTANICAL AND ECOLOGICAL CHARACTERISTICS
• FIRE ECOLOGY
• FIRE EFFECTS
• MANAGEMENT CONSIDERATIONS
• REFERENCES

INTRODUCTORY

AUTHORSHIP AND CITATION FEIS ABBREVIATION NRCS PLANT CODE COMMON NAMES TAXONOMY SYNONYMS LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
	Gary P. Fleming © DCR Virginia Natural Heritage Program

Hybrids: Broadleaf cattail hybridizes with both other North American cattail species, narrow-leaved cattail (T. angustifolia) and southern cattail (T. domingensis), where distributions overlap. Hybrid swarms of all 3 species have been identified in central California. T × glauca has been used to describe both broadleaf cattail × narrow-leaved cattail and broadleaf cattail × southern cattail hybrids [36,72,107,143,194,227]. In this review, hybrids are identified with both parent names.

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• HABITAT TYPES AND PLANT COMMUNITIES

Cattail hybrids are often identifiable by their distribution. Distributions of all 3 hybridizing cattail species overlap in only the east-central US coast and central California. Broadleaf cattail and southern cattail cooccur in a broad area of the southwestern United States and the extreme southern United States. Broadleaf cattail and narrow-leaved cattail distribution overlaps are more common and cover a broad area of the central and eastern United States and Canada. Cattail distribution maps are provided by Smith [194].

Often wetlands are dominated by one or just a few clonal species occurring in zones or concentric rings that are largely determined by flooding and water level tolerances [41,146,209]. Numerous wetland plant communities and habitat types are described, and broadleaf cattail is often recognized as a dominant species.

• broadleaf cattail floating wet mat communities in the northwestern part of Tanana Flats south of Fairbanks [174]

•  

• bulrush (Scirpus spp.)-cattail tule marshes common in the Central Valley [24]

• brackish hardstem bulrush (Schoenoplectus acutus var. acutus)-broadleaf cattail marshes common in San Francisco Bay [137]

• tule marshes along the Sacramento and San Joaquin rivers; marshes part of wet grasslands Forest and Range Ecosystem No. 41 [67]

• tule marsh potential natural vegetation type [115]

• tule marsh and cattail-sedge (Carex spp.) communities; variants of the wetland Rangeland Cover Type (Cheatham and Haller, cited in [2])

• softstem bulrush (S. tabernaemontani)-broadleaf cattail and broadleaf cattail wetlands throughout the state (references in [8])

• floating island type vegetation dominated by beggarticks (Bidens spp.), broadleaf cattail, and smartweed (Polygonum spp.) in Orange Lake in Alachua County [139]

• broadleaf cattail marshes and cattail associations near brackish water in southern Florida [41]

• broadleaf cattail habitat types along ponds and reservoirs at elevations of 2,980 to 5,740 feet (909-1,750 m) in the east [86]

• broadleaf cattail plant associations throughout the state [104]

• broadleaf cattail-softstem bulrush marshes in the Chilly Slough Wetland Conservation Area in Custer County [237]

• broadleaf cattail-tussock sedge (Carex stricta) communities along creek borders in Cook County's Bluff Spring Fen Nature Preserve [200]

• broadleaf cattail dominated marshes in the Gavin Bog and Prairie Nature Preserve in Lake County [204]

• softstem bulrush-cattail-bur-reed-rush (Sparganium-Juncus spp.) in swales and river system depressions; southern and narrow-leaved cattail possible, broadleaf cattail probable

• cattail-Olney threesquare (Scirpus americanus) in oxbows and low areas along creeks and streams; all cattail species possible

• cattail-scouringrush horsetail-sedge (Equisetum hyemale) on hillsides and bluffs in river valleys; broadleaf and/or narrow-leaved cattail may dominate [123]

• broadleaf cattail-California bulrush (S. californicus) associations in southeastern freshwater marshes [171]

• broadleaf cattail herblands in riparian areas [12]

• broadleaf cattail habitat types in northwestern Montana [18] and central and eastern Montana [90]

• annual wildrice (Zizania aquatica)-broadleaf cattail and broadleaf cattail-hardstem bulrush communities in Cherry County [209]

• broadleaf cattail-Olney threesquare and broadleaf cattail-rice cutgrass (Leersia oryzoides) community types in the upper and/or middle Rio Grande watersheds [52]

• broadleaf cattail-common threesquare (Schoenoplectus pungens var. pungens) communities widespread in Pecos and Rio Grande basins, likely throughout New Mexico

• softstem bulrush-broadleaf cattail vegetation throughout New Mexico [160]

• broadleaf cattail western herbaceous vegetation type within the cattail-bulrush (Schoenoplectus spp.) semipermanently flooded herbaceous alliance [165]

• tule marshes around large lakes; marshes part of wet grasslands forest and range ecosystem No. 41 [67]

• broadleaf cattail-rush-Jamaica swamp sawgrass (Cladium mariscus subsp. jamaicense) and rush-broadleaf cattail-Virginia saltmarsh mallow (Kosteletzkya virginica) freshwater marsh communities on Shackleford Bank [7]

• interdune pond communities (vegetation varies with water depth)

• broadleaf cattail-black rush (Juncus roemerianus) tidal freshwater marsh communities [184]

• cattail-bulrush hydric freshwater marsh associations on the Coastal Plain; broadleaf and narrow-leaved cattail common [226]

• broadleaf cattail herbaceous vegetation associations on floodplains, backswamps, and lake margins

• broadleaf cattail-powdery alligator-flag (Thalia dealbata) associations in marshes, ponds and on lake margins in eastern and central Oklahoma [100]

• broadleaf cattail palustrine emergent wetland vegetation types in Gettysburg National Military Park [236]

• hardstem bulrush-broadleaf cattail marshes around Utah Lake in Provo Bay and Powell's Slough [21]

• tule marshes on old beds of the Great Salt Lake; marshes part of wet grasslands Forest and Range Ecosystem No. 41 [67]

• hardstem bulrush-broadleaf cattail communities in ponds with 1- to 3-foot (0.3-0.9 m) deep water in steppe vegetation in the Columbia Basin [39,61]

• broadleaf cattail plant associations in eastern National Forests [113]

• broadleaf cattail-toad rush (Juncus bufonius) community types in narrow seeps on the Pumice Plains of Mount St Helens [43]

• broadleaf cattail-bog goldenrod/sphagnum (Solidago uliginosa/Sphagnum recurvum-S. imbricatum) communities in Laurel Run Bog

• broadleaf cattail/bristly dewberry/recurved sphagnum (Rubus hispidus/S. recurvum) communities in Cupp Run Swamp [217]

• broadleaf cattail cover types on wet or moist sites throughout the state [32]

• broadleaf cattail community types at low elevations of Utah and southeastern Idaho [176]

• semiaquatic wetlands of the rooted emergent class dominated by broadleaf cattail and hardstem bulrush in Idaho and western Montana [173]

• broadleaf cattail types in semipermanently and saturated palustrine wetlands in the prairie potholes of north-central United States and south-central Canada [106]

• broadleaf cattail/broadleaf arrowhead (Sagittaria latifolia) plant associations in Wyoming, Nebraska, and Colorado [105]

• broadleaf cattail marshes in Montana, Wyoming, Colorado, Nebraska, and New Mexico

• bulrush-cattail marshes in the northern Great Plains (Manitoba, Minnesota, Montana, and North and South Dakota)

• Olney threesquare-cattail communities in the central and southern Great Plains (Colorado, Kansas, Nebraska, New Mexico, Oklahoma, and Texas); broadleaf, narrow-leaved, and southern cattail possible

• cattail-American lotus (Nelumbo lutea) in Oklahoma and Texas; broadleaf and southern cattail common [186]

• softstem bulrush-broadleaf cattail reed swamp associations of large sloughs and lakes in central Alberta [130]

• broadleaf cattail marsh associations in the Cariboo Forest Region [196]

• broadleaf cattail vegetation types in marshes along the Ottawa River between Ontario and Quebec [42]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

GENERAL BOTANICAL CHARACTERISTICS RAUNKIAER LIFE FORM REGENERATION PROCESSES SITE CHARACTERISTICS SUCCESSIONAL STATUS SEASONAL DEVELOPMENT
	© Br. Alfred Brousseau, Saint Mary's College

Aboveground description: Broadleaf cattail is an aquatic or semiaquatic emergent perennial. Morphological and physiological ecotypes as well as hybrids make field identification difficult [82]. Plants are normally 3 to 10 feet (1-3 m) tall, reedlike, and extensively clonal [98,122,172]. Broadleaf cattail stems are stout, cylindrical, and unbranched [5,172]. Flowering stem length is typically equal to or somewhat longer than leaf length [143]. Leaves are thick, linear, flat, and measure 6 to 29 mm wide [5,98]. For plants from Michigan and the southeastern United States, the upper limit of leaf width is less, 15 mm [72,216]. Broadleaf cattail can be highly productive. In a single growing season, a plant grown from seed produced 34 aerial shoots that were 18 to 24 inches (46-61 cm) tall, 29 shoots that were 4 to 18 inches (10-46 cm) tall, 35 shoots measuring 2 to 4 inches (5-10 cm), and 104 lateral buds [238].

Numerous tiny, dense, felty flowers occur in a terminal spike that is 0.7 to 2 inches (1.8-5 cm) thick [46,110,143,172]. Male flowers make up the upper, narrower half of the spike and female flowers the lower, slightly wider half [82,108]. Over 1,000 flowers were counted in the staminate spike portion of one plant [98]. Male and female spikes are rarely separated. A small space of less than 0.6 inch (1.5 cm) may occur [82,108], but separations of up to 1.5 inches (4 cm) are also reported [82]. Some systematists recognize plants with separated male and female spikes as form ambigua [187,216]. Spikes are typically 6 times as long as they are thick [46,110,143,172]. Only female flowers are persistent [143,172]. Fertilized flowers produce single-seeded, nutlike achenes up to 1.5 mm long [122,227]. Long slender hairs at the base allow for wind and water transport of the "eventually dehiscent" seeds [99,172].

Belowground description: Floras from throughout broadleaf cattail's range describe its rhizomes as tough, stout, coarse, and extensive [36,98,99,172]. Rhizomes grow horizontally just below the soil surface [82]. In a review, lateral rhizomes were reportedly up to 28 inches (70 cm) long, with diameters of 0.2 to 1.2 inches (0.5-3 cm) [76]. Shallow fibrous roots are attached to the rhizomes [10,47]. Some indicate that broadleaf cattail produces 2 types of rhizomes; one that is "more superficial", thin, "feathery", and multibranched, and another that is deeper, thicker, and branching at the base [181]. Rhizome growth and depth are likely affected by substrate texture, moisture, and/or temperature. In the Skokie Marsh of Illinois, the thickest broadleaf cattail rhizomes were 3 to 3.9 inches (7.5-10 cm) deep [190]. From an alluvial basin in central Iowa, long, stout broadleaf cattail rhizomes occurred at 6- to 8-inch (15-20 cm) depths. Rhizomes were soft and spongy, with internal air spaces [96]. For more on vegetative growth, see Vegetative regeneration.

© 2003 George W. Hartwell

Broadleaf cattail × narrow-leaved cattail hybrids, often referred to as T × glauca, have been described as more "robust" and "competitively superior" than either parent in fluctuating water levels, although hybrids are infertile and reproduce only vegetatively [122,216]. Invasiveness of broadleaf cattail × narrow-leaved cattail is discussed briefly in Other Management Considerations [234].

Pollination: Female broadleaf cattail flowers mature before male flowers, making cross pollination possible or even likely [10,82].

Breeding system: Broadleaf cattail is monoecious [5,72]. Heterozygosity and polymorphism levels suggest extremely low genetic variation in broadleaf cattail stands, but genetic differentiation exists between stands; ramets within a site are more similar than those between sites. Researchers indicate that self pollination and clonal growth have increased the genetic homogeneity of stands while increasing genetic differences between sites. Increased genetic diversity of broadleaf cattail occurred at the most polluted sites along a transect from Louisville, Kentucky, to Circleville, Ohio. Researchers were not sure that pollution levels caused the genetic differences [109].

Seed production: When broadleaf cattail flowers, abundant seed production is possible. A review reports that typically more than half of pollinated flowers set seed [102]. In controlled conditions, broadleaf cattail grown from seed produced flowers early in the second year of growth. From 10 spikes that averaged 7 inches (18 cm) long, an average of over 222,000 seeds/spike was produced [238].

Many studies report poor flower production by broadleaf cattail. Sexual reproduction was rare in broadleaf cattail vegetation on the west side of Lawrence Lake in south-central Barry County, Michigan. Of 1,779 marked shoots studied over 3 years, only 3 flowered [45]. Degree of shading or depth of submergence may affect flowering. In broadleaf cattail monocultures in experimental ponds at the University of Arkansas, broadleaf cattail flowered only when roots were not submerged. Researchers suggested that self shading in dense stands may inhibit flowering [75]. In a manmade pond at Michigan's W. K. Kellogg Biological Station, the percentage of broadleaf cattail flowering in a single year in September varied with water level. Unsubmerged plants did not flower. When submerged under 6 inches (15 cm) of water, 33% flowered, and in 20 inches (50 cm) of water, 11% flowered [80]. Perhaps a water level influence on flowering is related to the nonflooded or barely flooded conditions necessary for successful seed germination. For additional information on this topic, see Germination.

Seed dispersal: Broadleaf cattail seeds are transported by wind, water, and substrate movement. Achenes have numerous long slender hairs at the base that allow fruits to float on water and blow in the wind [172], and some report that achenes split or burst when they contact water [92,98]. Broadleaf cattail produces abundant seeds [58]. A review describes the release of broadleaf cattail fruits from the spike. When fruits are dry, protective portions of the pistil shrivel and hairs on the achene spread. Spreading hairs produce pressure that bursts inflorescences and releases fruits into the air [102]. Wind dispersal distances of broadleaf cattail seeds collected in Hockessin, Delaware, were calculated from timed descents made indoors. Seed mass averaged 0.05 mg. Seeds fell at 0.13 m/s, and the estimated lateral distance traveled in a 10 km/h wind was 154 feet (46.9 m) [144].

Seeds are also dispersed through soil movement when mud clings to animals or people [47]. Seeds may also be transported with portions of broadleaf cattail clones that are torn by wind, water, ice, or animals [6]. Often seeds remain attached to spikes through the winter and are dispersed in the spring, and in some cases seeds fail to disperse. Apfelbaum [6] observed submerged cattail spikes with hundreds of nearby seedlings.

Seed banking: Broadleaf cattail produces a persistent seed bank. Although there are no reports of how long broadleaf cattail seed remains viable in the soil, its emergence from soils in late-seral forests that have long been unsuitable broadleaf cattail habitat suggests long-term persistence or long-distance dispersal. In Washington, a small percentage of field-collected broadleaf cattail seed germinated after being stored in a freshwater canal for 60 months [33]. While dense broadleaf cattail seedlings often emerged from sites where broadleaf cattail was important [126,210], there were exceptions [48,210]. Most of the studies below used the emergence method to determine the density of broadleaf cattail seed in the soil. This method requires, but rarely assures, that ideal broadleaf cattail germination conditions are provided.

Depth, season, and vegetation type: Broadleaf cattail emergence from soil collected in the Hamilton Marshes of New Jersey's Delaware River wetlands varied with season of collection, depth of burial, and vegetation type. Through field experiments, researchers determined that more than half of broadleaf cattail emergents came from persistent seed banks and not recent seed rain. The greatest number of broadleaf cattail seedlings (15,060/m²) emerged from the top 0.8 inch (2 cm) of soil collected in March from vegetation dominated by calamus (Acorus calamus) and broadleaf cattail. Soils collected in June from the same depth and vegetation type had 2,340/m² broadleaf cattail seedlings emerge. There were 5,060/m² and 2,150/m² broadleaf cattail seedlings in the top 0.8 inch of soils collected in March and June, respectively, in shrublands dominated by red maple (Acer rubrum), silky dogwood (Cornus amomum), and alder (Alnus spp.). Broadleaf cattail seedling emergence decreased with increasing soil depths, and no seedlings emerged from soil samples taken from depths greater than 5.9 inches (15 cm). More seedlings emerged on unprotected sites (53,000/m²) than on sites with seed rain excluded (38,700/m²). Although differences were not significant (P>0.05) [126], they suggest a persistent seed bank. For additional information on Leck and Simpson's [126] study, see Germination and Seedling establishment/growth.

High levels of seedlings or germinants are not always recovered from soils where broadleaf cattail is important. Few broadleaf cattails germinated from soils collected in eastern Tennessee mixed-deciduous forests. Soil samples collected in the spring from a site where broadleaf cattail frequency was 70% had 27±10 (SE) broadleaf cattail germinants. No seedlings germinated from summer-collected soils. From a site where broadleaf cattail frequency was 10%, there was only 1 germinant/m² from spring-collected soils [48].

Newly-colonized sites: Broadleaf cattail seed was recovered from new substrates resulting from the eruption of Mount St Helens. Seed banks develop soon after broadleaf cattail colonization [238]. In late September, soil samples were collected from newly-developed wetlands. No broadleaf cattail seedlings emerged from soil collected in wetlands where broadleaf cattail cover averaged 60%. In a wetland where broadleaf cattail cover averaged 90%, 126±145 (SD) seedlings/m² emerged from the top 2 inches (5 cm) of soil, and 1,088±1,609 seedlings/m² emerged from 2- to 4-inch (5-10 cm) depths. No broadleaf cattail seedlings emerged from soil samples that were not cold stratified. Researchers suspected that soil samples contained more broadleaf cattail seed than what emerged and that germination conditions were not ideal in the greenhouse [210].

Disturbed and late-seral forests: In south-central British Columbia, broadleaf cattail seeds emerged from undisturbed, 1-year-old "lightly burned", and 1-year-old clearcut Douglas-fir (Pseudotsuga menziesii) forests, although broadleaf cattail was not part of the existing vegetation. Broadleaf cattail seedlings emerged from soil samples collected from 5% of the quadrats in the undisturbed area, from 5% of lightly-burned quadrats, and from 2% of clearcut quadrats [195]. Few broadleaf cattail seedlings also emerged from the litter or top 0.8 inch (2 cm) of soil collected from 130- to 175-year-old mixed-conifer forests in eastern Oregon's Blue Mountains, although broadleaf cattail was not present in study plots or adjacent areas. Researchers suggested that broadleaf cattail seed survives in the soil until conditions are conducive to germination, allowing broadleaf cattail to rapidly occupy disturbed sites [202]. Low numbers of broadleaf cattail seedlings also emerged from soil collected in mature Douglas-fir and fir (Abies spp.) forests averaging 88 years or older in central Idaho [114].

Germination: Specific requirements necessary for broadleaf cattail germination are difficult to ascertain. Experimental studies indicate that light and warm temperatures are necessary for broadleaf cattail seed germination. However, studies involving flooding, leaf litter extract, and stratification effects on seed have produced conflicting results.

Broadleaf cattail seeds collected for 3 years from populations at 8,500 feet (2,600 m) elevation near Grand Lake, Colorado, failed to germinate. Researchers noted that broadleaf cattail had occupied the site for at least 12 years and that vegetative growth was vigorous [149]. Reasons for the variability in germination potential, requirements, and tolerances are unknown. Possibly broadleaf cattail seed viability is temporally, environmentally, ecotypically, and/or genotypically variable.

Cold stratification: As part of an extensive study of broadleaf cattail ecotypic population differences, McNaughton [147] found that broadleaf cattail seeds did not require a cold period before germinating, but the lowest temperature at which 50% germination occurred was lower in southern than northern populations. Broadleaf cattail seeds collected in soils from newly-developed wetlands on Mount St Helens only germinated in soils that were stratified (37 °F (3 °C)) for 12 weeks. No seedlings emerged in unstratified wetland soils [210].

Salinity: Choudhuri [27] found that broadleaf cattail germination percentages decreased with increasing salinity. No seeds germinated above 1 atm of osmotic pressure in natural salinity. For additional information on broadleaf cattail's tolerance of salt as a juvenile and adult plant, see Soil salinity.

Light, pH, temperature, oxygen, and ash: Broadleaf cattail seeds germinate best in warm temperatures and high light conditions [19,191,224]. Seeds germinate in acid, basic, or neutral pH conditions, and ash extracts have increased broadleaf cattail germination [178]. Reduced oxygen levels through the manipulation of gases in the air or through submersion have also increased broadleaf cattail germination success [191].

Broadleaf cattail seeds collected from North Bay Park on Ford Lake Reservoir shores in Washtenaw County, Michigan, germinated better in light than in dark conditions, better at 77 °F (25 °C) than at 68 °F (20 °C), and better when submerged than when exposed. Exposed seed germination, regardless of temperature or light exposure, ranged from 0.8% to 5%. Germination was best, 79.2%, when seeds were submerged, in the light, and kept at 77 °F (25 °C) [224].

Germination increased with increasing temperatures for broadleaf cattail seeds collected in May and October from Carlos Avery Wildlife Management Area, Minnesota. Maximum germination percentages occurred at the maximum temperature tested, 95 °F (35 °C). No seeds germinated at 50 °F (10 °C). Seeds exposed to cold temperatures before exposure to 95 °F (35 °C) temperatures had lower germination percentages than seeds kept at room temperature. Germination did not occur when seeds were exposed to low oxygen levels of 1.0 mg/L, and at least 10 hours of continuous red light was required for maximum germination. Researchers suggested that broadleaf cattail germination in the field would be most likely at the surface of saturated soils [19].

Optimum broadleaf cattail germination temperatures were 77 to 86 °F (25-30 °C), and seed crops responded differently to reduced and normal oxygen levels in studies conducted in Ontario. Germination rates were slower and percentages were reduced at 68 °F (20 °C) and 95 °F (35 °C). Seeds immersed in water germinated better than those kept moist. When moist seeds were exposed to reduced oxygen levels (2%), germination percentages equaled those of immersed seeds. Light exposure increased germination. The light intensity required for germination was low, but critical intensities varied with individual seeds. Seeds in full light had 99% germination. In containers wrapped with 3 layers of herbarium paper, germination was 97%, and when wrapped with tin foil, germination was 5%. Germination studies spanned many years, and differences were found among broadleaf cattail seed crops. Seeds collected in 4 of 7 years had increased germination in low oxygen environments, but 3 crops showed little to no apparent difference with oxygen levels changes [191].

Collection time affected the germination of broadleaf cattail seed collected in November and May in central Alberta. Less than 1% of fall-collected seeds germinated in light and distilled water, but 90% of seeds collected in May germinated. Germination in light (81-90%) was significantly higher than in the dark (15-73%, P<0.01). Germination in the dark increased when broadleaf cattail ash was added to distilled water. Germination was not affected by pH levels of 4, 7, or 12. Researchers suggested that high light levels and ash on burned sites may favor broadleaf cattail seed germination. Researchers also suggested that low humidity levels during seed storage may have affected viability [178], but spring collected seeds may have benefited from natural stratification.

Flooding: Many controlled experiments indicate that broadleaf cattail seeds germinate well in submerged or flooded conditions; however, field observations suggest regeneration from seed is restricted to nonflooded substrates [189]. Beule [16] indicated that field observations made in southeastern Wisconsin did not agree with laboratory reports of broadleaf seed germination in flooded conditions. Only once did Beule find broadleaf cattail seedlings in areas where germination may have occurred under shallow water. All other field observations suggested that germination was restricted to exposed substrates [16]. Differences between laboratory and field observations may relate to light requirements. The quality and murkiness of water in the field and water used in the laboratory were likely different, and associated vegetation cover in the field was not likely mimicked in laboratory studies.

Broadleaf cattail seeds collected near Huntley, Montana, germinated at very low percentages regardless of pH or temperature treatments. Only when seed coats were ruptured along the edge were high germination percentages achieved. Broadleaf cattail seeds with ruptured seed coats germinated under 30 inches (76 cm) of water and grew to the surface [238]. Broadleaf cattail germination percentages were greatest, about 43%, at 16 inches (40 cm) deep and lowest, approximately 16%, when unflooded at the Aquatic Environmental Research Facility in Lewisville, Texas. Germination percentages at 31-inch (80 cm) and 39-inch (100 cm) depths were about 25% [188].

Emergence of broadleaf cattail from reclaimed coal mine wetland soils in Perry County, Illinois, was better when samples were submerged in 0.8 inch (2 cm) of water than when kept moist [31]. Flooding soils collected from the Hamilton Marshes of New Jersey's Delaware River wetlands with 1.2 to 1.6 inches (3-4 cm) of water did not significantly affect broadleaf cattail germination (P>0.05). Researchers noted that broadleaf cattail emergence in the field was substantially less than in the greenhouse. Frequency of broadleaf cattail seedlings in the field ranged from 0 to 47% over a 2-year period in 2 vegetation types [126]. See Seed banking for more on seedling emergence results from Leck and Simpson's [126] study.

Nutrients and extracts: Some studies report differences in percent germination and germination rates when broadleaf cattail seeds are exposed to leaf litter extracts and nutrients. Germination rate increased but total percent germination was not different when seeds from the northern Florida Everglades were kept moist with Everglades water rather than distilled water. Everglades water with low, medium, and high levels of total phosphate was tested [198].

Germination was nearly a complete failure when seeds collected from broadleaf cattail marshes near Syracuse, New York, were kept moist with broadleaf cattail leaf extracts. In distilled water, broadleaf cattail germination was 90.8% after only 2 days. The researcher described the inhibition as "autotoxic feedback", a process by which seedling viability decreases with accumulations of toxic parent residues [148]. Germination of broadleaf cattail seeds from Washington County, Arkansas, was inhibited by 3% broadleaf cattail leaf extract concentrations, but inhibition was due to the development of water molds. There were no significant differences in germination of seeds planted in pots with (82%) or without (86%) adult broadleaf cattail (P>0.05). Light levels were high (81% full sun) and litter accumulations were low in the pots, which is not representative of a natural broadleaf cattail stand [74]. In natural broadleaf cattail marshes in Crawford County, Pennsylvania, light was less than 3% at the soil surface [23].

Broadleaf cattail hybrids: Increased sediments and flooding reduced broadleaf cattail × narrow-leaved cattail germination percentages. Germination of broadleaf cattail × narrow-leaved cattail seed collected from Larson Marsh in Story County, Iowa, decreased by 60% to 90% when 2 to 4 mm of sediment were added to the soil surface [218]. In the greenhouse, broadleaf cattail × narrow-leaved cattail and broadleaf cattail germination was 15% and 16%, respectively, in 16 inches (41 cm) of water. Germination was best in 1 inch (2.5 cm) of water, and germination percentages decreased with increasing water depths [225].

Seedling establishment/growth: Broadleaf cattail seedlings are extremely small when compared to seedlings of associated vegetation. Drawings of typical broadleaf cattail seedlings are provided by Leck and Simpson [125]. After germination, broadleaf cattail produces 2 to 4 small leaves and 2 to 6 floating leaves before producing erect leaves. Once shoots reach 14 to 18 inches (35-45 cm) tall, rhizome growth begins (Holm and others as cited in [155]). Flooding and sediments can affect seedling survivorship and growth.

Flooding: Some indicate that broadleaf cattail seedling establishment is most likely on nonflooded substrates [16,189]. However, Yeo [238] observed that nearly all broadleaf cattail seedlings in the field were submerged in early growth. At the Aquatic Environmental Research Facility in Lewisville, Texas, broadleaf cattail seedling biomass was measured 10 weeks after germination at 0- to 47-inch (120 cm) depths. Seedling biomass was greatest at 7.9 inches (20 cm) and least at 47 inches (120 cm). Unflooded conditions produced seedling biomass greater than flooding with 24 or more inches (>60 cm) [188].

Hybrid seedlings/growth: Broadleaf cattail × narrow-leaved cattail seedling survivorship was much lower in 1 cm of sediment than when no sediment was present, and younger seedlings were more sensitive to sedimentation than older (26-37 days) seedlings. Survival decreased when over half of the shoot was covered. Survivorship increased when older, larger seedlings received small amounts of sediment. Sediment loads of up to 1.6 inches (4 cm) did not affect adult plant densities [218].

Growth: Typical broadleaf cattail growth and development were summarized after 3 years of studies on the west side of Lawrence Lake in Michigan. There are 3 shoot emergence pulses/year; one occurs in the early spring, with spring shoots dying by late fall. In midsummer another pulse of growth occurs, and 70% to 80% of shoots die in the fall. The last growth pulse produces 80% to 90% of the shoots that will resume growth in spring [45]. The rhizome is the most long-lived broadleaf cattail structure; it survives up to 2 years (Westlake, cited in [45]). Broadleaf cattail growth and production can be affected by soil moisture regimes and associated species presence.

Broadleaf cattail biomass production and height growth were greatest for continuously flooded seedlings in a greenhouse experiment. Researchers collected broadleaf cattail seed in March from freshwater wetlands along the Mississippi River in Tennessee. Seedling biomass production and height growth were significantly lower in periodic drought conditions than in continuously flooded conditions (P<0.001) [131].

At the W. K. Kellogg Biological Station in Michigan, broadleaf cattail shoot density was 32% lower when growing with narrow-leaved cattail than when growing in a monoculture [81].

Vegetative regeneration: Rhizome growth is important to broadleaf cattail regeneration. Rhizome dispersal may occur when portions of a clone are separated by wind, water, ice, or animals [6]. Dispersal is also likely through tillage and substrate movement [47].

Broadleaf cattail is highly productive through clonal growth. Broadleaf cattail clones can occupy 58 m² two years after germination [77]. Holm and others [102] reported in a review that a single broadleaf cattail colony was 54 m² after 2 growing seasons and produced a total rhizome length of 1,600 feet (480 m). A plant grown from seed produced rhizomes that reached a diameter of 10 feet (3 m) in a single growing season [238]. Nearly monotypic broadleaf cattail stands are multiclonal [109]. Broadleaf cattail ramets typically die within a year. If the ramet flowered, it was very unlikely to live beyond 1 year. In undisturbed broadleaf cattail stands, ramets were not over 3 years old [78].

In many cases, vegetative regeneration predominates over sexual reproduction. Some indicate that seedlings rarely occur in established broadleaf cattail stands [78] and that rhizome production and growth are the primary methods for increasing stand size [44]. Researchers collected seed for 3 years from an established high-elevation broadleaf cattail stand near Grand Lake, Colorado. No seed germinated, but vegetative growth was described as vigorous [149]. A total of 1,765 broadleaf cattail shoots were monitored in the western marsh of Lawrence Lake in south-central Michigan over a 2-year period. There were no seedlings, and only 2 shoots flowered. Researchers summarized that "through vigorous vegetative growth", a dense, monotypic broadleaf cattail stand produced a "tightly packed advancing front of ramets" that successfully excluded other plants [44]. After a series of field experiments, researchers concluded that broadleaf cattail "is exploitive in its ability to clone rapidly and colonize available space, is able to capture light effectively because of its high allocation to leaves and high leaf surface area, and as a result has a low allocation to sexual reproduction" [79].

Climate: Broadleaf cattail's wide distribution implies a wide tolerance of climatic conditions. Broadleaf cattail occurs in tropical, subtropical, southern and northern temperate, humid coastal, and dry continental climates [76]. The following climate descriptions represent some widely different climates experienced in broadleaf cattail habitats in the northern and southernmost portions of its North American range.

Northern habitats: Scattered broadleaf cattail populations occur in central Alaska, where winter temperatures are as low as -31 °F (-34 °C) [76]. Ponds in open subarctic woodlands near Yellowknife, Northwest Territories, occur in subarctic-continental climates and remain frozen from late September to mid- or late May. Winters are long and cold; summers are cool and short with very long periods of sunlight. On 21 June, it is light for 20 hours [60]. Daily temperatures can vary widely in broadleaf cattail marshes of the Ottawa River between Ontario and Quebec. Daily average minimum and maximum temperatures for January are 5 and 23 °F (-15 and -5 °C) and for July are 54.5 and 77 °F (12.5 and 25 °C). An average of 120 days are frost free [42].

Southern or desert habitats: Broadleaf cattail occurs in Diamond Pond in the semiarid desert of Harney County, Oregon. Relative humidity is low, and evaporation is high. A typical growing season is 80 to 117 days. Annual rainfall averages 7.9 to 12 inches (200-300 mm), and snow levels average 33 inches/year. Daily and seasonal temperatures fluctuate widely. In nearby Burns, Oregon, temperature extremes of -33 °F (-36 °C) and 100 °F (39 °C) have been reported [229]. Broadleaf cattail is also found on the edges of Mitry Lake near Yuma, Arizona. This site received 3.5 inches (89 mm) of precipitation in 1963 and 1.5 inches (38 mm) in 1964. Minimum and maximum temperatures in 1964 were 31 °F (-0.5 °C) in January and 104 °F (40 °C) in July [145]. Broadleaf cattail and narrow-leaved cattail-California bulrush communities in southeastern Texas experience frequent droughts and occasional hurricanes. Typically, there are only 6 or so days of freezing temperatures. In Port Arthur, rainfall averages 51.8 inches (1,320 mm). Over a 20-year period, precipitation averages ranged from 31 to 66 inches (840-1,700 mm) [124]. In treeless wet prairies of southern Florida's Everglades region, broadleaf cattail occupies sites with high summer rainfall, high humidity, mild winters, and often a midwinter dry season [135].

Elevation: Broadleaf cattail occupies sites from sea level to 7,500 feet (2,300 m) throughout North America [58]. Elevations at more narrow geographical ranges are given below.

Soils: Broadleaf cattail tolerates many soil textures, nutrient levels, moisture regimes, and pH levels. Descriptions of soil types as well and flooding, drought, and salinity tolerances are described below.

Texture, pH, and fertility: Sand, silt, loam, and clay substrates are described in broadleaf cattail habitats. Acid to basic soils with low or high levels of nutrients are tolerated. In the Cariboo Forest Region of British Columbia, broadleaf cattail marshes occupy silty clay loam soils with seasonal or permanent water up to 39 inches (100 cm) deep [196]. The top 4 inches (10 cm) of wetland soils on the Pumice Plain of Mount St Helens, Washington, were predominantly silt (67.2-80.7%). Organic matter was low (<0.54%), and pH was 5.7 to 7.2 [210]. Broadleaf cattail in oxbow lakes along the Athabasca River in Alberta occupied basic sites with a pH of up to 9.2 and water depths up to 9.4 inches (24 cm) [132]. Germination of broadleaf cattail seed from central Alberta was not affected at pH levels 4, 7, or 12 [178]. In the Lake Agassiz Peatlands Natural Area of Minnesota, broadleaf cattail was indicative of weakly minerotrophic waters with calcium levels of 3 to10 ppm [97]. In Wisconsin, broadleaf cattail occurred in water with less than 50 ppm and in waters with more than 150 ppm of calcium carbonate [37]. Broadleaf cattail marshes of the Ottawa River between Ontario and Quebec occupy the most fertile temporarily flooded sites available [42]. Detailed descriptions of soil fertility in broadleaf cattail stands in Mississippi, Alabama, Georgia, and Florida are provided by Boyd and Hess [20]. In these areas, broadleaf cattail standing crop biomass was positively correlated with the concentration of phosphorus in the soil and water (r =0.69).

Floating mats: Sometimes broadleaf cattail occurs on floating mats with minimal soil development. In a freshwater marsh northeast of Sackville, New Brunswick, floating mats of broadleaf cattail × narrow-leaved cattail were 16 to 24 inches (40-60 cm) thick in water 31 to 39 inches (80-100 cm) deep. Developing small mats got their buoyancy from aerenchyma (tissue with air spaces) in the rhizomes, while older, thicker mats floated on trapped air bubbles produced during anaerobic decomposition [101]. In oxbow lakes along the Athabasca River in Alberta, broadleaf cattail occurred on floating mats in nutrient-poor lakes and was rooted on more nutrient-rich sites [132].

Soil salinity: Broadleaf cattail tolerates brackish waters; however, salinity tolerance may differ with developmental stage. Choudhuri [27] found that broadleaf cattail germination decreased with increasing salinity, and no seeds germinated above 1 atm of osmotic pressure. However, broadleaf cattail was "quite tolerant" of salinity as a seedling and as an established plant. In marshes of southeastern Louisiana, broadleaf cattail occurred at salt levels up to 1.13% [171]. In oxbow lakes along the Athabasca River of Alberta, broadleaf cattail occurred where salinity, measured as water conductivity, was 260 to 1,380 μs/cm [132]. A review of the vegetative characteristics of prairie marshes in western Canada reports that broadleaf cattail tolerates salinity levels of less than 10 mS/cm [189].

Flooding: Broadleaf cattail is tolerant of fluctuating water levels [189] and some flooding; however, death or colonization failure has occurred at flood levels as low as 25 inches (63 cm) [197], and stands grow in 3 feet (1 m) of water in wetlands of southeastern Alberta [111]. Flood tolerance has been associated with rhizome production, season, age, associated vegetation, and disturbances.

Broadleaf cattail grew in a maximum water depth of 27 inches (68 cm) in monoculture ponds created at the W. K. Kellogg Biological Station of Michigan and in a maximum depth of 24 inches (61 cm) in ponds with narrow-leaved cattail [81]. In the same ponds, broadleaf cattail grew at 31-inch (80 cm) depths in May but was restricted to less than 31-inch (80 cm) depths in September [77]. In experimental ponds at the University of Arkansas, broadleaf cattail died when water depths exceeded 37 inches (95 cm), and broadleaf cattail density was greatest at 8.7-inch (22 cm) depths. Broadleaf cattail height increased with increased water depths, but plants flowered only when roots were not submerged [75]. In a storage pool at the Montezuma National Wildlife Refuge in New York, broadleaf cattails less than 1 year old died when water levels were at or above 18 inches (46 cm). Two-year-old seedlings did not spread when water levels were 18 to 20 inches (46-51 cm), and at depths over 25 inches (63 cm), even well-established plants died [197].

In controlled field and greenhouse studies, broadleaf cattail and broadleaf cattail × narrow-leaved cattail rhizome production decreased in water levels of 12 inches (30 cm) or more [225]. In the 1st year of pond flooding in areas drawn down for 2 to 5 years in northwestern Minnesota's Agassiz National Wildlife Refuge, broadleaf cattail and broadleaf cattail × narrow-leaved cattail grew well at all water depths. After 3 years of flooding, broadleaf cattail death was substantial in 18 inches (46 cm) of water. Broadleaf cattail × narrow-leaved cattail growth and survival were good in up to 24 inches (61 cm) of water. By the 5th flooded year, broadleaf cattail was dead in all continuously flooded areas, but survival was good in temporarily flooded areas [93]. In the Crescent Pond south of Lake Manitoba, the density of broadleaf cattail × narrow-leaved cattail was greatest at 9.8-inch (25 cm) and 39-inch (100 cm) depths. Density was lowest at 33 inches (85 cm). Shoot height generally increased with increasing water depths, suggesting a plastic growth response [220].

Flooded conditions after cutting or burning often result in death. Abovewater plant material is necessary to tolerate flooding. When broadleaf cattail and broadleaf cattail × narrow-leaved cattail stems were cut below the water surface and kept flooded, both species died [225]. Broadleaf cattail rhizomes in a highly anaerobic environment in southern Michigan maintained high concentrations of oxygen as long as some portion of the plant remained above water. Plants cut below water 3 times in a growing season experienced almost complete belowground death. Researchers suggested that aerenchyma throughout the stems and rhizomes transported oxygen from the atmosphere to the rhizomes [183].

Drought tolerance: Broadleaf cattail is described as "fairly drought tolerant" [89]. In the Montezuma National Wildlife Refuge on the northern end of Cayuga Lake in New York, broadleaf cattail stands grew in 0.9 inch (2.2 cm) of standing water in June but were dry in July and August [57]. In marshes of Utah that were drained and kept dry for 2 years, there was 100% broadleaf cattail mortality [164]. The potential for broadleaf cattail regrowth with flooding was not discussed.

General descriptions: Most often, broadleaf cattail is described in early-seral communities. It is considered an early-seral species in bogs of Michigan [216]. It is a dominant emergent in early successional stages in low- and high-nutrient sites in oxbow lakes along the Athabasca River in Alberta [132]. The broadleaf cattail type is described as an "early successional wetland community" at low-elevation sites of southeastern Idaho and Utah [169]. In northwestern Montana, broadleaf cattail is described as a "pioneer species that tends to form a relatively stable type" [18].

Shade tolerance: Broadleaf cattail is shade intolerant [116], and full light conditions are most conducive to seed germination [19,191]. A review reported that shade levels of 40%, 60%, and 90% reduced broadleaf cattail growth by 60%, 80%, and 90%, respectively, when compared to full sun. Broadleaf cattail often died after 30 months in 90% shade [102]. Broadleaf cattail seedling height was lower in 33% full sun than in full sun conditions when grown from seed collected from North Bay Park in Michigan. After 84 days in the greenhouse, seedlings averaged a little over 24 inches (60 cm) under shade cloth and a little over 33 inches (85 cm) in full sun [224].

Primary succession: Broadleaf cattail appears early in the primary succession of open water and newly-deposited substrates resulting from avalanche and volcanic flows.

Hydrarch succession is the process by which shallow water species succeed and eventually replace deep water forms [226]. In open water succession, broadleaf cattail is typically present in the emergent vegetation stage following stages dominated by submerged leaf and floating leaf species. The submerged and floating leaf stages of hydrarch succession in floodplains, deltas, and oxbow lakes of Alberta are typically dominated by pondweed (Potamogeton spp.) and pond-lily (Nuphar spp.), respectively. Broadleaf cattail is common in the next stage. Eventually sites succeed to sedge meadows that are eventually colonized by cottonwoods (Populus spp.) and willows (Salix spp.) (Raup, cited in [159]),[130,214]. Very similar succession is described on Steeny Kill Lake in New Jersey [168], swamps of northwestern Minnesota [55], peatlands of Dane County, Wisconsin [63], and in Gulf Coast marshes of Louisiana and Texas [136], although important species within the submerged, floating leaf, and emergent stages are often different. Time frames for successional turnover were not discussed.

Debris flow succession: New substrates resulting from volcanic and avalanche debris flows may be colonized by broadleaf cattail as early as 1 year after deposition. On mud flows created from a slurry of rock, sand, mud, and organic material that melted with the Mount St Helens eruption, broadleaf cattail was present within 1 year of eruption [87]. Broadleaf cattail wetlands were also found on new pumice substrates north of the Mount St Helens crater 14 years after eruption [208]. On debris avalanche deposits along the North Fork Toutle River Valley caused by the same eruption, broadleaf cattail frequency was 5% three years after the eruption. Nine years after eruption, frequency had increased to19%. Twenty years after eruption, frequency had decreased to 2% [38].

Secondary succession: Broadleaf cattail rapidly colonizes moist to wet disturbed sites. Often broadleaf cattail is absent before disturbances but appears soon after canopy opening. Broadleaf cattail's persistent seed bank is likely important in the rapid colonization of disturbed sites.

General: In West Virginia wetland communities, broadleaf cattail was often associated with recent disturbance [59]. In bogs of northern lower Michigan, reed (Phragmites spp.)-cattail vegetation is common on sites "greatly disturbed" by fire or trampling. In Smith's Bog, broadleaf cattail was abundant in disturbed agricultural areas, but after farms were abandoned, broadleaf cattail dominance decreased [68]. Along the Saugus River in Wakefield, Massachusetts, the density and composition of disturbed broadleaf cattail marshes were not different from undisturbed marshes 1 year after power line construction (P>0.05) [167]. Broadleaf cattail colonized newly-deposited sand on Minnesota Point in Lake Superior within 1 year of dredging. Seedlings appeared 1 year after dredging, and in the next growing season, there were 9 broadleaf cattail seedlings/10 m² at a 10-foot (3 m) distance from the bay shore. Four years after dredging, the sand fill was dominated by a dense willow thicket and broadleaf cattail marsh [119].

Broadleaf cattail densities and population dynamics were compared in developing and maturing seral communities in Lawrence Lake area of Barry County, Michigan. Broadleaf cattail density was 10.4 ramets/m² in open marshes dominated by short-statured vegetation exposed to full sun and wind conditions. Density was greater, 21 ramets/m², in monotypic broadleaf cattail stands sheltered from wind. In adjacent woodlands dominated by red-osier dogwood (Cornus sericea subsp. sericea) and willows, broadleaf cattail density was much lower, 4.8 ramets/m². Broadleaf cattail growing-season mortality was lowest in open marsh and greatest in woodland communities. Researchers indicated that woodland ramet mortality was likely due to decreased light availability. In open marsh habitats, overwinter mortality was 2.5 to 4 times greater than in monotypic cattail stands and woodlands, respectively. Lack of protection from weather likely caused this discrepancy in overwinter mortality. Broadleaf cattail's presence in woodland communities was maintained by vegetative reproduction from the broadleaf cattail marsh [78].

Logging: In the Clay Belt of northeastern Ontario and western Quebec, broadleaf cattail occurred in ruts created by logging equipment in wetland black spruce (Picea mariana) stands. Forests were logged an average of 9.3 years before the study. Sites were mesotrophic, high in pH and nitrogen, and dominated by speckled alder (Alnus incana subsp. rugosa) and cottonwood [22].

Fire: Broadleaf cattail occurred in burned baldcypress (Taxodium distichum var. distichum)-dominated Pocomoke River Swamps on the eastern shore of Maryland. Broadleaf cattail clumps and monotypic stands occurred on shallow burned sites. Researchers noted that broadleaf cattail is a pioneer "fire weed" in postfire hydrarch succession [15]. Time since fire was not reported. Similar findings of broadleaf cattail in early postfire communities are presented in Fire Effects.

Grazing/trampling: Broadleaf cattail abundance can be reduced by common muskrat and livestock grazing. Common muskrats feed extensively on broadleaf cattail roots and rhizomes. Common muskrat populations can affect successional development and productivity in broadleaf cattail stands [37]. In a wetland north of Cambridge, Maryland, a researcher observed heavy broadleaf cattail feeding by common muskrats. Twelve years after heavy use, the broadleaf cattail marsh was converted to green arrow arum (Peltandra virginica)-dominated vegetation. Extensive common muskrat feeding converted solid broadleaf cattail rhizome mats to unconsolidated, anoxic, organic substrates, and water levels were lowered by 2 to 6 inches (5-15 cm) [66]. Additional information is available in Common muskrats.

Although broadleaf cattail is not considered particularly palatable to livestock, it may be consumed when water levels drop and/or other upland forage is unavailable. Grazing and trampling can reduce broadleaf cattail abundance. In northwestern Montana, heavy livestock use may convert broadleaf cattail communities to a Nebraska sedge (Carex nebrascensis) type [18]. In wetlands of southeastern Alberta, heavy cattle grazing and trampling killed nearly all broadleaf cattail seedlings on exposed mud. Trampling did not affect established stands. The researcher concluded that "broadleaf cattail as an established emergent is little affected by cattle, but as a moist soil stand it is often badly trampled" [111].

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

Fire regimes: Broadleaf cattail is restricted to moist or wet sites; however, some indicate that these habitats can burn frequently [40,65]. Fires are not considered frequent in all broadleaf cattail habitats, though. In alluvial communities of the southeastern Coastal Plain, broadleaf cattail occurs at the edge of oxbow lakes, where fire is not common [29]. It is likely that fire regimes in broadleaf cattail marshes and stands are dictated by surrounding upland vegetation. If nearby vegetation is highly flammable and conditions are dry, fire is likely in broadleaf cattail vegetation [65]. Fuels: Fires are not uncommon in broadleaf cattail habitats, and often fuel loads are more than adequate for fire spread. Robertson [179] reported that the fuel load/unit area in wetlands can be higher than that of uplands in the upper Midwest. Wetland fires may burn "hotter" and, given proper conditions, "faster" than upland sites. Fires in cattail marshes produce thick, black smoke, similar to that produced when tires burn (Rhode, personal communication in [179]).

© 2005 Louis M Landry, photo taken in Lac Boivin, Granby, Quebec

Pre- and early-settlement fires: Several studies report that Native people as well as early trappers and settlers burned wetland vegetation to improve travel, hunting success, and food availability. In northern Alberta, Native people concentrated burning in wetlands (Lewis, personal communication in [206]). The Kumeyaay of southern California reportedly burned cattail marshes at 3-year intervals (Shipeck as cited in [11]). In south-central Manitoba, delta marshes were intentionally burned by early trappers to improve travel, expose common muskrat lodges and coyote, fox, and American mink dens, and concentrate wildlife into unburned areas. Early settlers often burned Manitoba meadows to improve forage quality. Meadow fires often escaped and burned adjacent marshes. Burning often occurred in the first warm days of spring [219]. For more on the use of broadleaf cattail by Native people and early settlers, see Other Uses.

Fires in presettlement time were frequent in several broadleaf cattail habitats across its range. In the Central Valley of California, freshwater marsh vegetation grows linearly and was likely important to north-south fire spread during dry conditions. Freshwater marsh vegetation is very productive and provides abundant fuel [232]. In the Driftless Area peatlands of Tempealeau County, Wisconsin, where broadleaf cattail is a common emergent, pollen records indicate that fire frequency was high before European settlement [40]. Frost [65] suggests that marshes in North Carolina's Croatan National Forest burned "frequently" in fires originating in flammable adjacent upland vegetation. Likely the fire frequency would have been similar to that of surrounding mixed-pine (Pinus spp.) and pond pine (P. serotina) communities, which typically burned every 1 to 3 years in presettlement times.

Spontaneous combustion: A report of spontaneous combustion was reported in marshlands along the shore of Lake Pontchartrain near Mandeville, Louisiana. Witnesses watched a fire "apparently ignited spontaneously" on 4 August 1924 in a time of "unprecedented drought". Water levels were several feet below the soil surface, and temperatures in neighboring towns were 100 to 104 °F (38-40 °C). Additional investigations in the area revealed that at least 100 separate fires were burning along an 18-mile stretch of marsh and pine vegetation. Other possible ignition sources were ruled out because of accessibility and timing. Weather reports indicated that heating and ignition conditions necessary for spontaneous agricultural fires occurred that day near Lake Pontchartrain. Other naturalists in the area suggested that ignition may have come from a creeping ground fire [215].

Changes in fire frequency: Fire exclusion has likely decreased fire frequency in broadleaf cattail habitats. On the North Fork of the Flathead River drainage, Montana, wet sedge meadows, where broadleaf cattail occurs on newly-disturbed sites, did not burn from 1926 to 1988. Researchers indicated that this was the longest fire-free interval since 1600 [231].

Prescribed fire practices may also reduce fire frequency in broadleaf cattail vegetation from what likely occurred in presettlement times. Fire lines used to control wildfires and prescribed fires are often constructed in the upland-wetland transition areas, where broadleaf cattail is common. Prior to active fire management, fires may have burned through the wetlands [64].

Communities listed below are those where broadleaf cattail has the greatest potential as a persistent species. Fire regimes typical of broadleaf cattail stands are likely related to fire regimes in adjacent upland communities. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

Seedling establishment occurs on burned sites [63], but the seed source is unclear. In several fire studies, broadleaf cattail plants were not present before the fire but appeared soon after the fire on burned sites. Whether or not burned site colonization was primarily from dispersed seed, a persistent seed bank, or both sources is uncertain. Numerous studies without prefire or unburned comparisons make regeneration inferences difficult. Also see Seed banking and Seed dispersal.

Postfire seedling establishment: Controlled studies suggest that recently burned sites provide good broadleaf cattail germination habitat. Broadleaf cattail seed collected in central Alberta germinated best in light (P<0.01), and germination increased when seeds kept in the dark were watered with a mixture of broadleaf cattail ash and distilled water. Total seedling length also increased with ash treatments in the light and dark. Researchers indicated that high light and ash levels on burned sites may promote broadleaf cattail germination [178]. After a summer fire on peatlands of Dane County, Wisconsin, the frequency of mature broadleaf cattail plants and seedlings was 16% to 96% on 1-year-old burned sites. The fire burned to an average depth of 1 foot (0.3 m) into the peat layer. Frequency of seedlings was greatest near a ditch with an established broadleaf cattail population [63].

Severe and/or dry-season fires: Numerous fire studies in broadleaf cattail habitats described the fires as severe, but rarely was broadleaf cattail abundance lower on burned than unburned sites in early postfire years. Broadleaf cattail occurred in severely burned marsh meadows in the Oregon Coast Range. Sites burned at least 11 years earlier. Broadleaf cattail was absent from adjacent mature western hemlock (Tsuga heterophylla)-Douglas-fir forests (about 300 years old) [163]. Burned and unburned wetlands in Mono Lake Tufa State Reserve, California, were only visibly different in litter abundance 1 year after a fall prescribed fire. Strip fires burned in the fall when wind speeds were low. Fires produced near-black smoke and flame lengths of up to 10 feet (3 m) and were described as "high intensity". In the following April, broadleaf cattail was nearly 1.6 feet (0.5 m) tall on burned sites, and burned sites were green before unburned sites [11]. After the September Red Bench Fire that burned wet sedge meadows in Glacier National Park, broadleaf cattail had 0.5% cover in the 2nd postfire year but was not reported in the 1st or 3rd postfire years. The fire burned when conditions were extremely dry and often burned to mineral soil. Prefire data were not given, but unburned meadows were dominated by beaked sedge (Carex rostrata) with near exclusion of other species [231]. After severe fires in smooth cordgrass (Spartina alterniflora) and sawgrass (Cladium jamaicensis) swamps of southeastern Texas, cattail increased in the understory. After another severe winter fire 2 years later, when conditions were dry, cattail increased even more [124].

© 2005 Ross Orr. Fall prescribed fire in a cattail marsh in Washtenaw County, Michigan

Invasion after fire: Broadleaf cattail appeared after severe fire or fires during dry conditions in baldcypress and pondcypress habitats. Two sites in pondcypress domes near Gainesville, Florida, were severely burned in December. Broadleaf cattail occurred only on burned sites and was not present before the fires. Broadleaf cattail was very rare in undisturbed pondcypress domes. In the 1st postfire year, broadleaf cattail cover was 10% to 12%. Cover was greatest, 30% to 38%, in the 2nd postfire year. By the 5th postfire year, cover of broadleaf cattail was reduced to less than 5% in the domes [54]. In pondcypress swamps of southern Florida's Corkscrew Swamp Sanctuary, burned, logged, and logged and burned sites were compared. Logging occurred about 25 years earlier, and burning occurred in June, 7 to 8 years earlier in an excessively dry year. Broadleaf cattail was absent from logged sites, but average importance values ranged from 2 to 8.5 on logged and burned sites. On only burned sites, broadleaf cattail's importance value averaged 2.2. On burned and burned and logged sites, pondcypress was absent and coastal plain willow (Salix caroliniana) dominated the canopy [84]. Broadleaf cattail did not occur in baldcypress-dominated woodlands of the Pocomoke River Swamp on Maryland's eastern shore, but after a fire, broadleaf cattail clumps and monocultures appeared on shallow sites. Researchers considered broadleaf cattail a pioneer "fire weed" in postfire hydrarch succession [15]. Fire severity, fire season, and time since fire were not reported.

Broadleaf cattail × narrow-leaved cattail: On the Hog Lake impoundment of the Tintamarre Marsh northeast of Sackville, New Brunswick, summer prescribed burning in drained areas significantly reduced broadleaf cattail × narrow-leaved cattail cover, density, and height, but summer burning in flooded areas increased cover significantly from unburned reference areas (P<0.05). Spring, summer, and fall fires occurred in flooded and drained areas of the marsh. Broadleaf cattail × narrow-leaved cattail cover, density, and height were compared 3 years after fire. In most spring and fall burn treatments, burned and unburned sites were rarely different, except broadleaf cattail height was significantly greater on spring-burned flooded marsh than on unburned flooded marsh (P<0.05) [138].

Postfire flooding/grazing: While most studies indicate that broadleaf cattail is highly tolerant of fire, increasing depths of postfire flooding often produce increased broadleaf cattail mortality. Preferential grazing on burned sites may also decrease broadleaf cattail abundance. In the Howard Slough of Utah, a prescribed fire burned most broadleaf cattail stems to the ground. Fire season and severity were not reported. The slough was flooded with 1 to 18 inches (2.5-46 cm) of water immediately after the fire. Broadleaf cattail growth was "heavy" on intermittently flooded and shallow-water sites (<8 inches (20 cm) "soon" after the fire, but it was absent from deep-water sites (8-18 inches (20-46 cm)) [164]. In Lacreek National Wildlife Refuge in southwestern South Dakota, cattail cover (narrow-leaved and broadleaf) was about double on unburned than on burned sites flooded with 4 to 12 inches (10-30 cm) of water after fire (P<0.05). Prescribed fires burned 2 sites in the fall and 2 sites in the spring. Cattail cover was similar on 1-year-old burned sites, regardless of burn season (P>0.05). Sites were without standing water by summer. Fires burned when fuel moisture was 41% to 253%, soil moisture was 36.5% to 123%, temperatures were 66 to 79 °F (19-26 °C), and relative humidity levels were 30% to 37%. Fires removed almost all aboveground fuels but did not consume any organic soil. Postfire herbivory was not different between protected and unprotected sites, indicating that grazers did not preferentially feed on burned sites [182].

Average annual production of broadleaf cattail was much lower on burned and grazed than undisturbed sites 1 year after treatments in the Ogden Bay Waterfowl Management Area freshwater marsh in Utah. Waterfowl and common muskrats preferentially grazed burned sites. Drawdown began in April, and prescribed fires burned in early September. Windspeed averaged 10.3 miles (16.6 km)/h, dew point averaged 41 °F (5 °C), and the maximum daily temperature was 83 °F (28.5 °C) during the fire. Sites were flooded 1 week after burning. Broadleaf cattail regenerated entirely through sprouting; there were no seedlings on burned sites. Burned and burned and grazed sites had lower broadleaf cattail production than unburned and ungrazed sites [193].

FIRE MANAGEMENT CONSIDERATIONS:
Fire and postfire flooding have been used to reduce broadleaf cattail abundance. Several challenges commonly occur when burning wetland habitats. Fuel loads are often high, and fires may burn "hotter" and "faster" than upland vegetation. Construction of fire breaks and maneuverability may also pose challenges in wetlands habitats [179]. Smoke produced in burning wetlands has been compared to the thick, black smoke produced when burning tires (Meyer, personal communication in [179]).

In the Carlos Avery Wildlife Management Area, managers used fire to maintain open water in dense cattail stands. Water levels were drawn down, and cattail vegetation was burned in the summer. Sites were flooded for several weeks, and cattail was typically removed for at least 2 years (Rhode, personal communication in [179]). Beule [16] reported that spring fires, ignited on a sunny day in drained marshes following a winter with very little moisture, are most likely to produce cattail mortality. Winds directing smoke away from populated areas are important when burning cattail wetlands [16].

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

Livestock: In northwestern Montana, broadleaf cattail habitat types may receive heavy livestock use if water levels are low and/or upland forage is limited [18]. In wetlands of southeastern Alberta, heavy cattle grazing and trampling did not affect mature established stands [111].

Native ungulates: Deer and elk use of broadleaf cattail is noted in Montana and Washington. Deer may use broadleaf cattail stands for food and shade or hiding cover in northwestern Montana [18]. Elk feces from the Mount St Helens blast zone in southwestern Washington had an average broadleaf cattail content of 0.1% to 1.3% from June to September [152].

Small mammals: Broadleaf cattail provides nest habitat and food for a variety of small mammals. In the Great Plains, broadleaf cattail is used as a lining in small mammal nests [203]. Grass-cattail habitats of No Name Lake along the lower Colorado River in Arizona had the highest capture rate per unit effort for Arizona cotton rats. Direct use of broadleaf cattail by the Arizona cotton rat was not discussed [3]. A study of gray catbird nesting concentrated in Michigan revealed that white-footed mice took over and modified these nests for their own. Cattail down was found in some of these nests and apparently brought in by white-footed mice [166].

In North Carolina, South Carolina, Louisiana, and Texas, broadleaf cattail was an important nutria food source. In marshes of North Carolina's Hatteras Island, broadleaf cattail was a primary summer food for nutria. Broadleaf cattail was consumed less in the winter [154]. A review cited several sources that identified broadleaf cattail or other cattails as important foods for nutria in Texas, Louisiana, and South Carolina [112].

Common muskrats: Broadleaf cattail is a primary common muskrat food, and broadleaf cattail habitats are used extensively for nesting. In the prairie pothole region of the north-central United States and south-central Canada, cattails are considered a "staple" common muskrat food [62]. In marshlands south of Saskatoon, Saskatchewan, broadleaf cattail was preferred winter habitat based on availability and use (P<0.05) [153]. Cattail stems are combined with mud to construct lodges. When common muskrats populations are high, cattail can decrease with extensive use [37]. In the Corinna marsh area of central Maine, broadleaf cattail was an important year-round common muskrat food. Broadleaf cattail frequency on feeding platforms averaged 45% overall and 85% and 87% in fall and winter, respectively [205]. Broadleaf cattail occurred in 78.9% of common muskrat feed beds in an alkaline fen on Upper Chateaugay Lake in New York [117]. After heavy common muskrat feeding on broadleaf cattail in a wetland north of Cambridge, Maryland, broadleaf cattail vegetation was converted to a green arrow arum-dominated type. The solid broadleaf cattail rhizome mat changed to an unconsolidated, anoxic, organic substrate, and the marsh was lowered by 2 to 6 inches (5-15 cm) [66]. In Gulf Coast marshes of southern Louisiana and Texas, common muskrats feed on broadleaf cattail primarily when Olney threesquare is unavailable [136].

Birds: Waterfowl and other marsh birds throughout the United States and Canada use broadleaf cattail habitats extensively. Specific studies and research are organized geographically in the paragraphs below.

Western habitats: Red-necked grebes, whooping cranes, marsh wrens, blackbirds, sandhill cranes, rails, and ring-necked pheasants all utilize broadleaf cattail habitats. Researchers located 89 red-necked grebe nests in natural and man-made ponds in Yellowknife, Northwestern Territories. Broadleaf cattail was used as a construction material in 43% of nests, and 39% of nest were anchored to broadleaf cattail [60]. In Wood Buffalo National Park in western Canada, whooping cranes used softstem bulrush-broadleaf cattail communities as nesting habitat [207]. Wrens, blackbirds, and waterfowl utilized broadleaf cattail as habitat and food in the Pacific Northwest [172]. In Oregon's Malheur National Wildlife Refuge, sandhill crane nests were better concealed and had higher success rates in hardstem bulrush and broadleaf cattail than in broadfruit bur-reed (Sparganium eurycarpum) or meadows. Nest rate success in broadleaf cattail vegetation was 58.8% [134]. Broadleaf cattail/hardstem bulrush marshes along the Colorado River between the Arizona/Nevada and US/Mexico borders were utilized by insectivorous and granivorous wading, water, and shorebirds [4]. In northwestern Montana, the broadleaf cattail habitat type is critical nesting and roosting habitat for red-winged and yellow-headed blackbirds [18]. Hen ring-necked pheasants on the Snake River Plain in southern Idaho used wetlands dominated by broadleaf cattail or willow for loafing, day escape, roosting, and thermal night winter cover more than expected based on availability (P<0.01) [129]. In Colorado, broadleaf cattail habitats were important nesting cover and habitat for soras, Virginia rails, blackbirds, and marsh wrens [83,221].

Central/eastern habitats: Canada geese, snow geese, mallards, sandhill cranes, marsh wrens, and blackbirds utilize broadleaf cattail habitats throughout central and eastern US and Canada habitats. In Marshy Point, Manitoba, Canada geese preferred broadleaf cattail to hardstem bulrush vegetation for nesting. Preferred nesting habitats were determined from nest occurrence and availability of the vegetation type. Over a 3-year period, 133 nests were found in broadleaf cattail. Nest predation was greater in bulrush, 24%, than in broadleaf cattail, 9% [34]. In the Delta Marsh of south-central Manitoba, breaking up continuous broadleaf cattail stands and increasing open water increased habitat use by nesting mallards. Use and production of dabbling ducks was maximized in the hemimarsh phase of development, when cover of water and emergent vegetation were nearly equal [162]. Sandhill crane nests were found in a wetland in Kennebec County, Maine. Nests were surrounded by broadleaf cattail and were constructed entirely of the previous season's broadleaf cattail stems and leaves [151]. In Kansas, broadleaf cattail vegetation is shelter and nest cover for red-winged blackbirds, yellow-headed blackbirds, and marsh wrens [10]. In Gulf Coast marshes of Louisiana and Texas, snow geese feed on broadleaf cattail roots and rhizomes in the winter when tides are low and Olney threesquare is unavailable. In a cold winter, snow geese "devoured several square miles of cattail marsh" [136].

Palatability/nutritional value: Nutritional value of broadleaf cattail was reported for southwestern Washington, northern Texas, and northern New York. Dry matter digestibility of vegetative broadleaf cattail collected in the summer from the Mount St Helens blast zone was 52%. Digestibility was 59.2% when plants were in flower. Vegetative plants were 2.95% nitrogen, and nitrogen was 2.93% in flowering plants [152]. The nutritive quality of broadleaf cattail was evaluated in the spring from control, grazed, and spring burned wetlands in Castro County, Texas. Differences between treatments were rarely significant. Broadleaf cattail protein ranged from 9.8% to 15%, ash was 10.3% to 13.4%, cellulose was 25.9% to 32.9%, and lignin was 13.4% to 19.5% [192]. Lacki and others [117] provide nutrient content and digestibility of broadleaf cattail collected in summer on the Upper Chateaugay Lake, New York.

Cover value: Broadleaf cattail stands are important cover for waterfowl, songbirds, and other wildlife species throughout its range [10,18,99,203]. Species specific information is available in Importance to Livestock and Wildlife.

In western Montana, laboratory and field experiments revealed that broadleaf cattail aerated soils and may have facilitated neighboring plant growth [25]. Planting broadleaf cattail on degraded sites may not be necessary, since this species rapidly colonizes disturbed sites through seed dispersal or germination from a persistent seed bank. See Seed banking, Seed dispersal, and Secondary succession for details.

OTHER USES:
Native Americans and early European settlers utilized broadleaf cattail as a food source, a construction material, and a medicine. Broadleaf cattail has numerous uses and has been referred to as the "supermarket of the swamps" [185]. Liptay [133] indicated that tribal wars were waged over control of broadleaf cattail marshes.

Food source: Broadleaf cattail is entirely edible, and Native Americans utilized broadleaf cattail year-round. Newly emerged sprouts were eaten as a green vegetable in the spring. Flower stalks were boiled and eaten like corn on the cob. Broadleaf cattail pollen, which has a nutty flavor and is high in protein, was added to other flours. Rhizomes were dug and eaten in the fall and winter. They were cooked or dried, pounded, and used in flour [1,10,133]. Comparisons of the nutrient values of broadleaf cattail, rice, and potatoes revealed that broadleaf cattail shoots and rhizomes contained much more calcium, iron, and potassium than potatoes or rice [133]. Broadleaf cattail was utilized as a food source by the Kawaiisu of south-central California [239], the Cahuilla of southern California [13], the Apache and other southwestern Natives [26], Native people along the Atlantic Coast [51], the Menominee of northern Wisconsin, the Ojibwe in Michigan [185], and likely many others.

Construction/ornamental/ceremonial material: Broadleaf cattail had a tremendous number of household and ceremonial uses. Leaves were used to make mats, dolls, baskets, and shelters. The hairs of fruits were used as sound-proofing material, insulation, pillow and lifejacket stuffing, and as tinder for fire starting. A paste made from rhizomes was used to caulk leaky canoes [10,133]. Early European settlers used the hollow broadleaf cattail stems to make candles [50]. Native people of the Pacific Northwest wove leaves into bedding, kneeling mats, capes, hats, blankets, and bags. Seed fluff was used to dress wounds and to stuff pillows, mattresses, and diapers [172]. Fluff material from seeds was used by the Menominee of northern Wisconsin and the Ojibwe of Michigan to insulate boots and jackets [185]. The same material was used by tribes of the Missouri River region as diaper material, pillow filling, and cradle board padding. Broadleaf cattail stem pieces were used as ceremonial objects by the Omaha and Ponca people [70]. Natives of southern California used broadleaf cattail leaves to construct houses as well as bedding [13,239]. Navaho people wore bracelets and necklaces made of broadleaf cattail leaves during the Male Shooting Chant ceremony, and dancers were dusted with broadleaf cattail pollen in other ceremonies [53]. The northern Cheyenne of Montana incorporated broadleaf cattail leaves into their Sun Dance [95].

Medicinal use: Broadleaf cattail was most commonly used as a wound dressing. Rhizomes were ground into a salve for wounds [133]. The Cahuilla of southern California used broadleaf cattail rhizomes to stop bleeding [13]. The northern Cheyenne of Montana made a tea of roots and leaf bases to treat stomach cramps [95]. The Sioux mixed "downy" cattail fruits with coyote fat to treat smallpox sores [94]. Tribes of the Missouri River region used fluff from broadleaf cattail seeds to make burn dressings [70]. For more on the past, current, and potential future uses of broadleaf cattail and other cattail species, see Morton [158].

Broadleaf cattail and broadleaf cattail × narrow-leaved cattail invasions: In some areas, broadleaf cattail has increased at the expense of other native species. In Harewood Marsh of Jefferson County, West Virginia, hydrologic changes and increased nutrient inputs may have facilitated broadleaf cattail increases, which may threaten rare species' persistence [49]. Nutrient enrichment also likely facilitated broadleaf cattail encroachment of sawgrass into the northern and central Everglades [35]. Urban runoff that increased nitrogen and phosphorus inputs coincided with broadleaf cattail × narrow-leaved cattail increases in Gardner Marsh in the University of Wisconsin-Madison Arboretum. Experiments revealed that broadleaf cattail × narrow-leaved cattail aboveground biomass, ramet density, and height were greater in fertilized than in control plots [234].

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