1. Azolla Lamarck in Lamarck et al., Encycl. 1: 343. 1783.
[Greek azo, to dry, and ollyo, to kill, alluding to death from drought]
Thomas A. Lumpkin
Roots 3--5 cm. Stems prostrate, 1--3 cm, or nearly erect, 3--5 cm, hairs absent. Leaves with 1(--2)-celled hairs on upper surface of upper lobe. Sporocarps in pairs. Megasporocarp megaspore with 3 floats. Microsporocarp masses entirely covered with arrowlike barbs. x = 22.
Species ca. 7 (3 in the flora): tropical to temperate regions.
Azolla is divided into sect. Azolla and sect. Rhizosperma (Meyen) Mettenius, which are sometimes recognized as subgenera. New World species belong to sect. Azolla and differ from sect. Rhizosperma by having 3 floats per megasporocarp (fig. 25.1), subdichotomous branching, and straight barbs on microsporangial masses (rather than 9 floats per megasporocarp, pinnate branching, and needlelike hairs or hairs absent on microsporangial masses). The genus was more diverse in past geologic ages: 30 species are known from the Cretaceous (J. W. Hall 1974).
The species of Azolla are difficult taxonomically because (1) about 80% of the specimens lack sori, which are necessary for identification, and (2) the characteristics needed to identify the species are difficult to observe. A scanning electron microscope is needed to see sculpturing of the megaspores, and a light microscope is needed to see the number of cells per hair on the upper leaf lobe. (These hairs are best seen in profile on mature leaves; at least 40× magnification is needed.)
Previous workers emphasized the number of septae (internal partitions) in the barbs on the microsporangial masses as a primary differentiating characteristic among species of Azolla (e.g., H. K. Svensen 1944). This character is not constant, however, either within a species or within an individual (R. K. Godfrey et al. 1961; L. V. Hill and B. Gopal 1967; K. Seto and T. Nasu 1975). Because nearly all floristic work in North America since the 1940s has been based on Svenson's synoptic treatment, the identity of most specimens is questionable, and therefore ranges are imprecisely known.
The maps for this treatment are tentative. For the most part, I have noted occurrences only in those states or regions from which specimens have been identified using characteristics given in this key. Literature that attributes a particular species of Azolla to a particular state or province must be questioned because the specimens were presumably identified using the inconsistent glochidial characteristics given by H. K. Svenson (1944). More work is needed to determine the distribution of Azolla species in North America.
Agriculturally, Azolla is famous for its symbiosis with the nitrogen-fixing Anabaena azollae Strasburger, a cyanobacterium (blue-green alga) found at the stem apices, beneath indusia, and in cavities of the upper leaf lobes. Because the plants fix nitrogen, they are often used as a green fertilizer or mixed with livestock feed as a nutritional supplement. Azolla pinnata has been cultivated for many centuries in rice paddies of northern Vietnam and southeastern China, where it acts as a fertilizer after it decomposes.
Azolla is the most frequently studied genus of ferns in the world because of its economic importance. The three North American species are naturalized in Europe and South Africa, and they have been introduced into Hawaii for horticulture and into Asia for agriculture. All species have been studied for agricultural uses in rice-producing areas.
Azolla is usually found in stagnant or slow-moving water of ponds, lakes, marshes, swamps, and streams. Plants turn reddish when under stress, such as from poor nutrition, salinity, or high temperatures. Sporulation needs further investigation.
Hall, J. W. 1974. Cretaceous Salviniaceae. Ann. Missouri Bot. Gard. 61: 354--367. Lumpkin, T. A. and D. L. Plucknett. 1982b. Botany and ecology. In: T. A. Lumpkin and D. L. Plucknett, eds. 1982. Azolla as a Green Manure: Use and Management in Crop Production. Boulder. Pp. 15--38. Moore, A. W. 1969. Azolla: Biology and agronomic significance. Bot. Rev. (Lancaster) 35: 17--35. Svenson, H. K. 1944. The New World species of Azolla. Amer. Fern J. 34: 69--84. Van Hove, C., T. de Waha Baillonville, H. F. Diara, P. Godard, Y. Mai Kodomi, and N. Sanginga. 1987. Azolla collection and selection. In: International Rice Research Institute. 1987. Azolla Utilization. Proceedings of the Workshop on Azolla Use. Fuzhou, Fujian, China, 31 March--5 April 1985. Los Banos, Laguna, Philippines. Pp. 77--87. Zimmerman, W. J., T. A. Lumpkin, and I. Watanabe. 1989. Classification of Azolla spp., section Azolla. Euphytica 43: 223--232.