213. Typhaceae A. L. Jussieu
S. Galen Smith
Herbs, perennial, of fresh to slightly brackish wetlands, often emergent, rhizomatous, caulescent in flower, smooth, glabrous. Leaves basal and cauline, 2-ranked, mostly ascending; sheaths open, margins overlapping, clear, summit tapered into blade or auriculate; blades twisted into loose helix, narrowly linear-attenuate, apex acute, aerenchyma prominent. Inflorescences 1, terminal, erect, equaled or exceeded by cauline leaves, cylindric, spikelike (hereafter "spikes"); staminate spike flowers deciduous but axis generally persistent; staminate spike distal to pistillate spike; young spikes subtended by early-deciduous bracts resembling reduced foliage leaves, 1 bract subtending pistillate spike, 1 bract subtending and several within staminate spike; ,staminate spike flowers deciduous but axis generally persistent; staminate axis with numerous simple or branched scales arising among flowers; pistillate axis with numerous projections ("compound pedicels"), evident on denuded fruiting spike, each bearing several flowers; in some species flowers subtended by slender bracteoles. Flowers unisexual, staminate and pistillate on same plants, numerous, densely packed in unisexual spikes, minute, wind-pollinated (stigmas receptive several days before pollen is shed); perianth probably represented by staminate scales and by hairs on stipes of pistillate flowers. Staminate flowers stipitate; stamens 1--several, filaments connate; anthers basifixed, connective distally extended. Pistillate flowers hypogynous, stipitate (stipe bearing numerous straight hairs, developing after flowering, acting in wind dispersal of fruits); pistils 1, 1-carpellate; ovariesy 1-locular; placentation apical; ovules 1; styles 1, unbranched; stigmas 1, whitish or green, drying brown, 1-sided, smooth; agamous flowers numerous (ovaries modified after flowering as carpodia). Fruits follicles, fusiform; pericarp clear, hyalinetransparent, splitting longitudinally in water to release seed. Seeds: endosperm starchy, oily; embryo cylindric.
Genus Genera 1, species ca. 8--13 (3 in the flora): boreal to tropical regions worldwide.
The extensive literature on morphology and taxonomy of Typhaceae has been recently reviewed by (U. Müller-Doblies and D. Müller-Doblies (1977); R. M. T. Dahlgren and H. T. Clifford (1982); R. M. T. Dahlgren et al. (19853); and J. W. Thieret and J. O. Luken (1996). The inflorescence is probably reduced from a compound structure.
Sparganium and Typha are very similar and perhaps should be placed in one family, as summarized by J. W. Thieret and J. O. Luken (1996): T (J. W. Thieret and J. O. Luken 1996). Other authors (e.g., D. Müller-Doblies 1970; U. Müller-Doblies and D. Müller-Doblies 1977; W. Schultze-Motel 1980) placed Sparganium in the Typhaceae.
Pre-Englerian [authors] ... placed Typha and Sparganium together in a single family, the Typhaceae. [H. G. A.] Engler (1886) put these genera in separate families, thus starting a tradition that has been followed by almost all subsequent authors until recently, when [D.] Müller-Doblies (1970) re-examined the relationships of the genera and concluded that "the five different characters by which Engler justified the family Sparganiaceae are wrong or, in two cases, without any significance ... The few remaining but very obvious differences may be explained to a large extend [sic] by an adaptation of Typha to anemochory [wind-dispersal of propagules]...."
The phylogenetic relationships of the Typhales with other families remain controversial, and it seems best to treat the taxon as an isolated order of uncertain relationships pending further research. Various authors have placed the Typhales close to or within the Pandanales, Arales, Poales, Liliales, Pontederiales, or Philydrales or in the Commeliniflorae generally close to the Cyperales and Juncales (J. W. Thieret and J. O. Luken 1996).
Briggs, B. G. and L. A. S. Johnson. 1968. The status and relationships of the Australian species of Typha. Contr. New South Wales National. Herbarium. 4: (2): 57--78. Dahlgren, R. and H. T. Clifford. 1982. The monocotyledons: a comparative study. 378 pp. New York. Dahlgren, R., H. T. Clifford, and P. F. Yeo. 1985. The families of the Monocotyledons. 520 pp. Berlin. Dugle, J. R. and T. P. Copps. 1972. Pollen characteristics of Manitoba cattails. Canadian Field Naturalist 86: 33--40. Eckardt, T. 1964. Pandanales. In: H. Melchior, ed. 1964. A. Engler's Syllabus der Pflanzenfamilien…, ed. 12. 2 vols. Berlin. Vol. 2, pp. 598--602. Engler, H. G. A. 1886. Ueber die Familie der Typhaceen. Bot. Centralbl. 25: 127. Fassett, N. C. and B. M. Calhoun. 1952. Introgression between Typha latifolia and T. angustifolia. Evolution 6: 367--—379. Finlayson, C. M., J. Roberts, A. J. Chick, and P. J. M. Sale. 1983. The biology of Australian weeds. II. Typha domingensis Pers. and Typha orientalis Presl. The J. Austral. Inst. Agric. Sci. 1983: 3--10. Grace, J. B. 1988. The effect of nutrient additions on mixtures of Typha latifolia L. and Typha domingensis Pers. along a water-depth gradient. Aquatic Bot. 31: 83--92. Grace, J. B. and R. G. Wetzel. 1982. Niche differentiation between two rhizomatous plant species: Typha latifolia and Typha angustifolia. Canad. J. Bot. 60: 46--57. Grace, J. B. and J. S. Harrison. 1986. The biology of Canadian Weeds. 73. Typha latifolia L., Typha angustifolia L. and Typha glauca Godr. Canad. J. Pl. Sci. 66: 361--379. Harris, S. W. and W. H. Marshall. 1963. Ecology of water-level manipulations on a northern marsh. Ecology 44: 331--343. Hotchkiss, N. and H. L. Dozier. 1949. Taxonomy and distribution of North. American cat-tails. Amer. Midl. Naturalist 41: 237--254. Kaul, R. B. 1974. Ontogeny of foliar diaphragms in Typha latifolia. Amer. J. Bot. 61: 318--323. Kronfeld, E. M. 1889. Monographie der Gattung Typha Tourn. (Typhinae Agdh., Typhaceae Schur-Engl.). Verh. Zool.-Bot. Ges. Wien 39: 89--192. Little, E. C. S. 1979. Handbook of utilization of aquatic plants. FAO Fisheries Tech. Pap. 187. Rome.. McNaughton, S. J. 1966. Ecotype function in the Typha community-type. Ecol. Monogr. 36: 297--325. Melchior, H. 1964. Pandanales. A. Engler's Syllabus der Pflanzenfamilien ed. 12. 2: 598--602. [incl. Typhaceae] Morton, J. F. 1975. Cattails (Typha spp.)-----weed problem or potential crop? Econ. Bot. 29: 7--29. Müller-Doblies, D. 1970. ÜÜber die Verwandschaft von Typha und Sparganium im Infloreszence Infloreszenz- und Blütenbau. Bot. Jahrb. Syst. 89: 451--562. figs. 1--224. Müller-Doblies, U. and D. Müller-Doblies. 1977. Typhaceae. In: G. Hegi, Illus. Fl. Mitteleuropa, ed. 3. II. 1(4), pp. 275--317. Berlin: G. Hegi et al., 1964+. Illustrierte Flora von Mitteleuropa, ed. 3. 5+ vols. in 8+. Berlin and Hamburg. Vol. 2, part 1(4), pp. 275—317. Ramey, V. 1981. Typha--—not just another weed. Aquaphyte. Newsletter of the IPPC Aquatic Weed Program of the University of Florida 1(1): 1: 1--2. Schultz-Motel, W. 1980. Typhaceae. In: H.J. Conert et al., eds. 1967--1980. Gustav Hegi's Illustrierte Flora von Mittel-europa. Pteridophyta, Spermatophyta. Band II. Angiospermae: Monocotyledones 2. Teil 1. Edition 3. Pp. 18--317. Berlin.. Smith, S. G. 1967. Experimental and natural hybrids in North American Typha (Typhaceae). Amer. Midl. Naturalist 78: 257--287. Smith, S. G. 1987. Typha: Its taxonomy and the ecological significance of hybrids. Arch. Hydrobiol., Beih. [Stuttgart]. 27: 129--138. Thieret, J. W. and J. O. Luken. 1996. The Typhaceae in the southeastern United States. Harvard Pap. Bot. 8: 27--56. Tompkins, T. M. and J. Taylor. 1983. Hybridization in Typha in Genesee County, Michigan. The Michigan Bot. 22: 127--131.