Introduction
When I first started researching this article, I did not know what I was getting into regarding the taxonomic problems of the Coontie. All I wanted to do was show everyone the different forms of Zamia that inhabit my home state of Florida. The more I dug, the more I found out that most people cannot agree with what this plant, or these plants, should be called. Although I will get into some of the details later, I am not the person to sort out and define what species these plants belong to. Instead of having a "focus on" article on Zamia integrifolia, or whatever the name should be, I felt that I was safe with "The Coontie of Florida" and let the taxonomists battle with the names.

History
There have been tribes of native people in Florida for the last 10,000 years. As new tribes came down from the main part of North America, these tribes changed. It is hard to say how long these plants have been used for food, but the first people known to eat this plant are the Calusa and Timucua Indians. In certain areas there were vast colonies of these plants, mainly in south Florida near Miami and Ft. Lauderdale. There is a reference to the mass quantities of plants around New River in Ft. Lauderdale; the native Indians called this place "Coonte Hatchee".

When the Seminoles moved into Florida in the mid-18th century, they picked up on this very important food source. In fact, "Coontie" is one of the names the Seminoles had for this plant and it roughly means "flour root". The Indians would cut up pieces of the stems and pound them out into a powder as much as possible. They would then wash this in water several times and then let the starch sink to the bottom. The paste was taken and fermented, and then dried to a powder. When the white men came to Florida they also used the stems for food. Their name for this plant was "Arrow Root".

Around 1845, several factories sprung up all over south Florida to produce starch from the Coontie. One of the mills along the Miami River is said to have processed 10-15 tons of product per day at peak production. In south Florida, a natural population would grow very slow. It can take 30 years to grow a plant that might weigh five pounds. These factories produced starch until 1925. Between the starch factories and the building in south Florida, only small remnants of these vast populations remain. I grew up in this area and I am afraid to say with all the building going on, there may only be a few isolated plants left.

Distribution
There are several forms of Coontie in Florida. There is a very thin-leaf form that grows on the eastern coast in St. Lucie County, in southern, central Florida. This plant is distributed all the way to the southern tip of Florida and into the Keys. The plant picks up again on the western coast in Everglades National Park; from there it runs north all the way up the coast to Taylor County, near the panhandle. There are old references that mention these plants growing halfway into the panhandle, but I do not know anyone who has seen them this far to the west. In most cases, these plants live in sandy soils in pine hammocks, although further to the North, the plants grow under turkey oaks as well. In Miami, and along some parts of the western coast, plants can be seen imbedded in coral rock; these plants show that Coontie can be grown in very alkaline soils and have a great tolerance to salt. All along the western coast, small plants can be seen growing on small outlying islands. The plants that grow in coral rock grow very slowly and are usually not very big. The plants that grow in sand can get much bigger.

Figure 1 shows a plant growing at the Montgomery Botanical Center, in an area that has been kept natural. Many of the plants that grow on rock or in semi-wet areas will have their apexes exposed. When I went to the habitat in Steinhatchee, a very xeric area, the tops of the tubers were at least thirty centimeters from the surface. Even though all these plants are called "the thin-leaf form," there is a lot of variation in certain areas. One unique plant, which has the thinnest leaflet of all the forms, occurs from the western side of Gainesville to Fanning Springs. The leaflets are also longer on this plant than on the other thin-leaf forms. (It reminds me a lot of Zamia portoricensis as far as leaflet size.) Figure 2 shows one of these plants in Fanning Springs. I had to put a white background in the picture just so the plant could be seen on film.

Figure 1. The south Florida Coontie in a natural stand at the Montgomery Botanical Center.	Figure 2. Coontie from Fanning Springs. This has the thinnest leaflets of any Coontie that I have ever seen.

In certain areas of the western coast, plants that have leaflets of moderate width can be seen. These plants grow in pockets and do not have large distributions. There is a lot of controversy over whether many of these populations are natural or whether these plants were brought in and cultivated by the Indians. It is interesting to note that many of these populations are alongside rivers, growing in sand. Figure 3 shows the distribution of the thin-leaf forms. Even though I agree with this map for the most part, I have heard of plants growing south and north of the shaded areas on the western coast.

Figure 3. Distribution of the thin-leaf form in Florida. Map prepared by Daniel B. Ward and Kent D. Perkins, who called this plant Zamia floridana.

The " wide-leaf " forms grow mainly in the northeastern part of Florida. The southern tip of its distribution starts in Brevard County and extends north into an area close to Jacksonville and west into Marion and Alachua Counties. This plant grows mainly in sand and usually grows under pines and oaks. Once again, in certain areas, plants can be seen growing where saltwater can flood for short periods of time. These plants will usually grow to be much larger, and I think they are much nicer looking for ornamental use.

Figure 4 shows a plant growing in Ocala National Forest. Again, within this area there are different looking plants. The most noteworthy plant has been called the "Palatka giant" by local people. This is truly an incredible plant. It grows more upright than most of the other forms; the leaflets are wider, the cones are larger, and the leaves have been said to reach 2.3 meters in length. Figure 5 shows a cultivated plant of the "Palatka giant" form growing in Gainesville. Figure 6 shows the distribution of the wide leaf form.

Figure 4. Plants growing in habitat in Ocala National Forest.	Figure 5. Russell Adams standing beside a "Palatka giant" plant that is in cultivation in Gainesville.

Figure 6. Distribution of the wide-leaf form in Florida. Map prepared by Daniel B. Ward, who called this plant Zamia umbrosa.

Taxonomy
In 1763, Linnaeus described Zamia pumila as coming from the Dominican Republic, Puerto Rico, and Cuba. This was the first Zamia to be described. James Eckenwalder feels that the Zamias from the Caribbean, including the plants from Florida, should all be Zamia pumila. There are many people who agree with this view on the Florida plants.

In 1789, William Aiton described Zamia integrifolia from a plant found in the Halifax River region--within the distribution of the wide-leaf form. According to Dan Ward, who works for the University of Florida in the Botany Department, the description of Zamia integrifolia is invalid. He says that in a certain area of the text, it is written that Z. integrifolia = Z. pumila, making these names synonymous and voiding the description. I could not find a copy of the original description to check this out for myself.

The next description of a Florida Zamia was in 1868, by Alphonse De Candolle, and was named Zamia floridana. This was one of the thin-leaf forms from an area north of Tampa. Then, in 1921, John Small described Zamia umbrosa, the wide-leaf form from around the Saint Johns watershed.

So what does this all mean? If you believe Eckenwalder, all the plants from Florida are Z. pumila. If not, then they are Z. integrifolia. If this latter speicies is invalid, then Z. floridana should be what they are called. If you believe that there are two separate species in Florida, the thin-leaf and the wide-leaf plants, then the thin-leaf form should be called Z. floridana and the wide-leaf form should be called Z. umbrosa. To confuse matters further, there are some people that feel there are more than two species in Florida. I have observed considerable variation in leaflet shape, cone size, and cone color on the different forms in Florida. The big question is "What is a large enough difference between two plants to make them separate species?"

Description
Stem
The stem of a single-headed plant can be 10 cm in diameter. Many times after a plant has produced a cone, the apex will split into two separate heads. After years of this, a stem can get to be very large. Also, large plants in the ground will push new tubers from the main stem, ending up in clumps up to 2 m in diameter. Stems are almost always subterranean unless the plants are growing on coral rock.

Leaves
Leaves can vary from 30 cm in length for the Fanning Springs plant to more than 2 m in length for the "Palatka giant". Leaflets vary in width from 2 mm for the Fanning Springs plant to 3 cm for the "Palatka giant". Most of the leaflets on the thin-leaf forms will curl up and twist, whereas most of the wide-leaf forms stay flat.

I have noticed that the leaflets of these plants will flatten out more if they are grown in the shade, whereas if you put them in full sun the leaflets will reduce in width and curl up sometimes. This appears to be a way to decrease the amount of moisture lost from the plant, by reducing the amount of surface area facing the sun. When my cultivated plants are pushing new leaves I will water them more. These plants will flatten out more than the same plants that I water less often. The typical habitats of the wide-leaf forms usually are shaded more than the habitats of the thin-leaf forms. This may have been the way that these plants have evolved over time. Figure 7 shows some of the leaflets from the different plants I have talked about.

Figure 7. Leaflets taken from some of the different forms in Florida.

Cones
Male cones can be 16 cm or more in length; they will usually be around 3 cm in diameter. Depending on the form, the cones can be black, brown, or even a dark orange. Figure 8 shows newly emerging male cones on a wide-leaf form.

Figure 8. Male cones emerging on a cultivated wide-leaf form.

Female cones can be only 6 cm long for a dwarf type from the west coast, to almost 30 cm long for a "Palatka giant" that is full of seeds. The width of the cone on the dwarf type can be as narrow as 4 cm, and the "Palatka giant" plant can have cones around 14 cm in diameter when they are filled with seeds. The colors of the female cones match the colors of the male cones. There are a lot of minor differences in the female cones from different areas. The nipple on top of the cone will vary from tall to almost flat. The female cone scales can vary as to the pattern and size. Usually, cones emerge around August and are receptive in January. Figure 9 shows an old cone and a couple newly-emerging cones on a wide-leaf form.

Figure 9. Older female cone with seeds and the new emerging female cones at the bottom.

Animal relationships
There are two beetles that pollinate the Zamias in Florida: Pharaxonotha zamiae and Rhopalotria slossoni. They will feed on the starch-rich male cones and are attracted to the female cones from what appears to be a change in color and temperature. I have not noticed any particular odor from the female cones but it is likely that these cones do emit some sort of odor not obvious to humans. The larvae of the Atala butterfly feeds on the leaves of the Coontie in south Florida, where the larvae of the Echo moth will eat the leaves of Coontie in north Florida.

There are many animals that aid in seed dispersal. Mockingbirds, grackles and blue jays seem to be the best dispersal agents. Rats and mice will move seeds to a lesser extent, but they will eat most of them. When the seed coat starts getting soft around April or May, many of these animals will carry off the seeds. Pill bugs will clean the seeds that are left around the mother plant. It is interesting to note that when the seeds drop off the plant, there is a waiting time before the seeds are ready to germinate. Right about the time that these animals start moving the seeds is when they should start germinating.

Cultivation
Most of the habitats have sandy soils. Thus, when growing these plants they should have good drainage. Most of the habitats are considerably shaded. Therefore, plants look their best when grown in partial shade; however, when grown in full sun they have the extra energy to produce more cones and leaves. Fertilizer should be applied on a regular basis, but the most significant time for producing leaves is in late spring. Cones emerge around August, so a high nitrogen fertilizer applied around June should help produce more cones. This is at least the timing in Florida. It would be interesting to see how these times change in the different parts of the world.

Acknowledgements
I would like to thank Daniel Ward for a lot of information on the taxonomy of the Coontie, as well as letting me use his maps of the distribution of the two forms in Florida. I would also like to thank Knut Norstog and Bart Schutzman for making me aware of the taxonomic problems. Also, I would like to thank Frank Brandt and Russell Adams for showing me around their areas to look at plants. And finally, thanks to Terrence Walters, Executive Director of The Montgomery Botanical Center, for letting me photograph plants in a habitat situation.

Bibliography
Eckenwalder, J. 1980. Dispersal of the West Indian cycad Zamia pumila L. Biotropica (12).
Small J. 1921. Seminole bread - The coonti. Journal of The New York Botanical Garden (22).
Tang, W. Florida's native cycad: Zamia pumila. Fairchild Tropical Garden Bulletin Jan. 1987.
Ward, D. B. (editor). Rare and Endangered Biota of Florida, Vol. 5.