Report SFRC-83/05 

The Vegetation of 

Long Pine Key, 

Everglades National Park 

Everglades National Park, South Florida Research Center, P.O. Box-279, Homestead, Florida 33030 
The Vegetation of Long Pine Key, 

Everglades National Park 

Report SFRC-83/05 

Ingrid Olmsted, William B. Robertson, Jr., JilJ Johnson, 

and Or on L. Bass, Jr. 

National Park Service 

South Florida Research Center 

Everglades National Park 

Homestead, Florida 33030 

August 1983 
Olmsted, Ingrid, William B. Robertson, Jr., Jill Johnson, and Oron L. Bass, Jr. 1983. 

The Vegetation of Long Pine Key, Everglades National Park. South Florida 

Research Center Report SFRC-83/05. 64 p. 
LIST OF TABLES 

LIST OF FIGURES 

LIST OF APPENDICES . 

ABSTRACT • 

INTRODUCTION. 

T ABLE OF CONTENTS 

CHAPTER 1 

• iii 

• iv 

v 

1 

3 

----HIST OR Y • 4 

~REVIOUS WORK 9 

METHODS . 11 

Vegetation Map • 11 

Numbers and Names of Hammocks • • 11 

Elevation Survey, Soils, and Hydrologic Parameters • • 12 

Quantitative Vegetation Analysis along Elevation Transect . • 12 

Hammock Fire History • 12 

Distribution of Shrub Stratum Plants in Long Pine Key Pinelands. • 13 

RESULTS. . 13 

Vegetation Map • 13 

Pinelands • • 14 

Pineland with Tall Hardwood Understory. • 15 

Tropical Hardwood Hammocks. • 15 

Hardwood Scrub Without Pine Overstory • • 15 

Hardwoods, Successional in Disturbed Areas • • 16 

Muhlenbergia Prairies • 16 

Prairie with Scattered Scrub • 17 

Prairie Species on Former Agricultural Land. • 17 

Cypress Prairie • • 17 

Cypress Forest • 17 

Willowheads • • 18 

Open Water • • 18 

Hole-in-the-Donut • 18 

Numbers and names of Hammocks • • 18 

Elevation Survey, Soils, and Hydrologic Parameters • . 19 

Quantitative Vegetation Analysis along Elevation Transect . • 19 

Hammock Fire History • 22 

Distribution of Shrub-stratum Plants in Long Pine Key Pinelands • 22 

SUMMARY • 25 
CHI\PTER l 

/PINELAND SHRUB UNDERSTORY: COMPI\RISON OF LONG PINE 

/ KEY WITH OTHER SOUTHERN FLORlI)l\ PINE FOREST~ 0 0 0 0 

METHODS 

RESULTS 0 

Regional Distribution and Diver!>ity of :-'hruh-straturn Plants 

Lower Florida Keys 0 

Long Pine Key 

Southern Biscayne 

Northern Biscayne 0 

Big Cypress 0 

Density Characteristics of thc Pincland :-'hrub Str.llurn 0 

DISCUSSION 0 

ACKNOWLEDGEMENTS 

LITERATURE CITED 

Ii 

o 26 

26 

o 27 

o 27 

o 2~ 

1\ 

o 11 

o 12 

on 

o 12 

13 

o ,4 

o 15 
LIST OF TABLES 

Table Page 

1. Ground water levels (em) in pineland wells · 41 

2. Summary of Vegetation Analysis . · 42 

3. 2 Mean cover percentage (per m ) and frequency of woody, 

forb, and grarninoid species in each vegetation type. · 43 

4. Frequency an~ percent cover of certain tree and grass 

species (10 m plots) indicating distribution in high 

and low elevation pineland sites · 46 

5. Fire impact status and inferred fire history of hammocks, 

by year, as determined from aerial photography and 

helicopter flights in 1981 · 48 

6. Distr ibution of 95 taxa of shrub-layer plants in pine 

forest areas of southern Florida · 51 

7. Summary: shrub stratum diversity and similarity in 

southern Florida pine forests · 54 

8. Density of shrub-stratum plants in southern Florida 

pine forests . · 54 
Figure 

l. 

2. 

3. 

LIST OF FIGURES 

Elevation transect from Osteen to Redd Hammock • 

Ordination of three vegetation types • 

Ordination of pineland stands • 

iv 

Page 

• 55 

• 56 

• 57 
LIST OF APPENDICES 

Appendix Page 

I . Numbers and names of Long Pine Key hammocks • 58 

II. Species list • 60 

v 
f 

ABSTRACT 

This report summarizes current knowledge of the vegetation of Long Pine Key, an 

extensive limestone region in the southern Everglades that constitutes the principal 

upland of Everglades National Park. 

Historical review revealed that Long Pine Key was disovered about 1840, but its 

location and dimensions were not firmly established until Krome's survey of 1902-

03. Since the area became accessible by road, farming (c. 1916-1976, initially in 

prairies, later also in uplands), lumbering of pine (c. 1936-1947), and prescribed 

burning of pine forests by the National Park Service (l958-present) have been the 

principal human activities that affected vegetation. 

Initial botanical exploration of Long Pine Key (mainly by Small beginning in 1909) 

showed that the upland flora was predominantly West Indian and contained a 

number of endemic taxa. Ecological studies have centered on the role of fire in 

successional relations between pine forests and tropical hardwood forests, the 

occurrence of rare and endemic plants in the area, and the invasion of abandoned 

farmlands by aggressive exotics, especially Schinus terebinthifolius. 

A vegetation map (scale 1:7800) was prepared from recent aerial photogr~phs to 

delineate the distribution of 12 plant cover types in an area of 7 x 15 km including 

most of Long Pine Key. Principal types of native vegetation were pine forests, 

tropical hardwood forests (hammocks) and Muhlenbergia prairies. 

Sampling along a 2 km transect provided data for quantitative description of the 

three principal plant communities in relation to site elevation, soil depth, and 

height of the water table. In all, 218 plant species were recorded of which 167 

occurred in pine forest, 114 in prairies, and 51 in hammocks. 

A definitive listing of the names of 120 Long Pine Key hammocks, originally named 

and numbered by tree snail collectors, was prepared to clarify previously confused 

nomenclature. 

Study of aerial photography for the period 1940-1980 showed that 90 percent of the 

Long Pine Key hammocks suffered detectable fire damage during this period, 

chiefly in 1945 through 1960, and that the course of recovery after fire varied 

widely between sites. Despite the effects of frequent fire, hammocks tended to 

retain their original size and shape over the period. 

The shrub understory of Long Pine Key pine forests was found to include a total of 

61 taxa 75 percent of which were of West Indian origin. As many as 29 species 

- occurred on plots .025 ha in extent and any of 19 species was locally dominant in 

the shrub understory. Between-site variation in shrub-stratum composition and 

diversity appeared to be related to long-term differences in fire effects and 

possibly to relict occurrence of several coastal plants in western areas of Long 

Pine Key. 
2 

Data for rockland pine forests throughout southern Florida showed a total shrub 

understory flora of about 95 taxa. The most diverse shrub stratum, about 60 taxa, 

occurred in the Lower Flor ida Keys and Long Pine Key areas decreasing to 40 in 

northern parts of the Biscayne Pineland of southeastern Florida and to fewer than 

30 in pine forests of Big Cypress National Preserve. Diversity was diminished 

primarily by the loss of West Indian species as one proceeded north in the mainland 

pine forests. A few shrubs characteristic of pine flatwoods of the southeastern 

United States occurred at the more northerly sites where limestone was overlain by 

acid sand. The shrub stratum of pine forests in the northern Bahamas appeared 

from floristic literature to be similar to that of southern Florida but considerably 

more diverse. 

• 
3 

INTRODUCTION 

Long Pine Key is the collective name applied to the rocky islands, surrounded by 

marsh, which extend west and southwest from Taylor Slough for about 25 km into 

the southern interior of Everglades National Park and form the southern boundary 

of the Everglades basin. Long Pine Key has a maximum elevation of about 5 m msl 

and its total extent of approximately 8000 ha constitutes the prinCipal upland area 

of Everglades National Park. The rock islands are exposures of a mid-Pleistocene 

oolitic limestone, the Miami Oolite, and represent an extension of the Atlantic 

coastal ridge into the interior of southern Florida (Hoffmeister 1974). The 

limestone formation was extensively modified by solution and erosion during low 

sea stands of the later Pleistocene and is cavernous at depth with the exposed 

portions extremely rough and pitted. Pioneer botanist, J. K. Small, gave perhaps 

the most graphic brief description of the substrate in stating (Small 1909:53), 

" ••• the surface consists mostly of holes." Fire-maintained forests of south 

Florida slash pine (Pinus elliottii var. densa) cover most of Long Pine Key, 

enclosing many small (seldom as large as 10 hal stands of broad-leaved forest, 

termed hammocks. Tropical taxa predominate in the upland flora somewhat 

justifying Small's (l916a: 39) characterization of the area as, " ••• a portion of the 

West Indies isolated on the Florida peninsuIf,.II Much more extensive pine-forested 

limestone uplands (approximately 400 km ) of similar character (but generally 

more elevated and less broken by erosion) formerly extended northeastward from 

Taylor Slough as far as the Miami River. However, except for a few small tracts 

preserved as Dade County Parks, urban and agricultural development has virtually 

eliminated native plant cover over this entire area. The pine rock lands of the 

Lower Florida Keys, also reduced by development with few areas assured of 

protection, differ considerably in aspect and floristics. Long Pine Key has, thus, 

become the principal remnant of a vegetation complex unique in the United States 

and the only area in which a number of tropical and endemic taxa are likely to 

survive in our flora. 

This publication is divided into two major chapters. In the first chapter, we review 

the history of exploration and previous botanical work, present a detailed map of 

plant communities in a major part of the area, and discuss various aspects of the 

Long Pine Key flora and vegetation. In particular, the distribution of tropical 

hardwood hammocks and their fire history was an important aspect of this study. 

Quantitative plant analysis was undertaken along an elevation transect through 

pine forest, hammock, and glades. The elevation survey gives information about 

the relationship between plant communities and the environment. The second 

chapter treats the understory shrub flora of Long Pine Key in relationship to other 

south Florida pine lands. We hope to establish a base from which future botanical 

and ecological studies of the area may proceed without the need of extensive 

historical investigation, and also, perhaps, to suggest lines which such studies might 

profitably pursue. 
CHAPTER 1 

HISTORY 

4 

At least as ear ly as the latter part of the 18th century it had been reported that 

pine-forested rocklands of a peculiar and forbidding character existed in far 

southern Florida. Thus, B. Romans (1962: 11) wrote of the soils in Florida pine 

forests: 

"In east Florida, in the southern parts, this kind of land is often very rocky, 

but especially from the latitude 25:50 southward to the point where it is a 

solid rock of a kind of limestone covered with innumerable small, loose and 

sharp stones, everywhere." 

Again (op. cit.: 192), describing the lands south of his Rattones River (d., Boca 

Raton): 

" ••. the remainder of the land is a heap of stones and rocks, very sharp, and 

little water to be found, there being only a few ponds, and these dry in a dry 

season. The only growth here is shrubby pine. II 

As far as it goes, Romans' description applies reasonably well to Long Pine Key, 

but it undoubtedly was drawn from areas immediately south of Miami, where pine 

rockland fronted on Biscayne Bay behind a narrow screen of mangroves and 

hammock and near anchorages commonly used by sailing ships. Romans' map of 

Florida and other maps as late as the early years of the Seminole Wars (Williams 

1837, Mackay and Blake 1839) suggest that the westward extension of the coastal 

rock ridge into the interior remained unknown. 

In the late 1830's, small military groups, moving Indian-fashion in dugout canoes, 

began to range widely in the southern Everglades and Big Cypress in search of the 

last Seminole camps (Griswold 1945). One result of this activity was the historical 

discovery of Long Pine Key. A careful search of military archives would be needed 

to identify the discovery positively, but one of the early visits (if not, in fact, the 

first) was that recorded in the diary of Army Surgeon J. R. Motte (Motte 1963). 

From a camp on Biscayne Bay near the present location of Cutler, Motte's party 

moved inland via a transverse glade and, thence, southwest to the end of the 

pinelands where "across a mile-wide prairie" they spied an Indian camp and fought 

a brief, inconclusive skirmish. The date was April 24, 1838, and a sketch map of 

the itinerary (op. cit.:221) locates the farthest point of advance on the eastern end 

of "Long Key." Motte'S account of the march through the rocklands (op. cit.:232) 

was rendered in rather purple prose, but anyone who has worked in the area would 

not find it altogether unfamiliar. 

" ••. we pursued our way through a pine-barren, the ground being formed of 

coral rocks jutting out in sharp points like oysterbeds, which caused us great 

suffer ing by cutting through our boots and lacerating our feet at every 

step. .. We suffered also very much for want of water, not a drop even of 

that which was stagnant was to be met with in this parched-up region. • •• It 
5 

was certainly the most dreary and pandemonium-like region I ever visited; 

nothing but barren wastes where no grateful verdure quickened, and no 

generous plant took root-where the only herbage to be found was stinted, 

and the shrubbery was bare, where the hot steaming atmosphere constantly 

quivered over the parched and cracked land--without shade--without water--

it was intolerable-excruciating. • •• But there was neither brook, nor bird, 

nor any living thing except snakes to be met with. About one o'clock P.M., 

we emerged from this rocky pine-barren ••• " 

The account is especially notable for its plain portrayal of the fact that very 

severe spring drought occurred in the area on occasion long before there had been 

any human manipulation of water. We shall endeavor to show that the rockland 

vegetation (which may have burned shortly before, although Motte does not say so) 

is not as devoid of interest as he found it. 

The sketch showing "Long Key," said to have been "reproduced from the Mackay 

and Blake map 1840" (op. cit.:221), depicts it as a narrow, crescentic reef about 

40 km long and trending northeast-southwest more or less parallel to the coast. 

This representation is repeated on eight other military and U.S. Coast survey maps 

of the 1840's and 1850's that we examined. Several of these, such as the wellknown Ives Map (Ives 1856), also show three military posts (Fort Henry, Camp 

Hunter, and Fort Wescott) in the general area of Long Pine Key. Perhaps these 

date from the March 1842 foray (Fort Dallas (i.e. Miami) to Long Pine Key to Shark 

River) of the command of a Lieutenant J. B. Marchand (Tebeau 1968). In any 

event, the later military expeditions do not seem to have added appreciably to 

geographical knowledge of Long Pine Key. 

C. W. Tebeau (op. cit.:76) relates that a Federal surveyor named Jack Jackson "ran 

the township lines in the Paradise Key (Royal Palm) area in 1847," but there seems 

to be no other information on this event. G. W. Cornwell and K. Atkins (1975:34) 

quote Tebeau without citing him in this connection, and they are surely in error in 

stating "his (i.e. Jackson's) old surveyor's trail made overland travel to Paradise 

Key possible, if slow, and an increasing number of scientists visited the area." 

Perhaps the mistake resulted from the fact that W. J. Krome, more than 50 years 

later, gave the name Camp Jackson to one of his advance survey points. In any 

case, no observers who left a record, scientific or otherwise, are known to have 

visited Paradise Key, Long Pine Key, or anywhere else in that area until after 

1880. The maps and reports of the Seminole War era seem to have been archived 

and, generally, forgotten. Many hunters and backwoodsmen in southern Florida 

doubtlessly knew of Long Pine Key and traversed (or avoided) it, but the area was 

unknown to the wor ld at large. Near the end of the 19th century, H. L. Willoughby 

(1898:124) could write with tolerable accuracy: 

"From the most easterly point reached on this day, I sighted the edge of Long 

Key and got a bearing on it. The existence of this large island at the 

southern end of the Everglades has been guessed at by white men, who have 

seen it from the edge of the pine timber bordering the Atlantic coast; but 

there is no accurate knowledge of its dimensions, and many have tried vainly 

to reach it." 
6 

All uncertainty about the location and size of Long Pine Key should have ended in 

1902~1-903 when Krome surveyed the area from Florida City to Cape Sable in the 

interest of a possible extension of the Florida East Coast Railroad. Notably, in the 

one pubJished item that came out of this remarkable episode, Krome himself 

(1904:453) counted as a major result of the survey the thorough mapping of "Long 

Key, a pine island of some 18000 acres, lying in the Everglades, the very existence 

of which was previously doubted •. " Krome (1903) prepared a detailed draft map 

of the uplands extending from the approximate site of present-day Homestead to 

the southwestern extremity of Long Pine Key and the rough trails to his camps at 

the eastern edge of Taylor Slough became the major overland access routes into 

the wilderness south of Perrine and Cutler for ear ly settlers and itinerant biologists 

(Smal1 1916a). The trail to Ca mp Jackson, located near the present eastern 

boundary of, Everglades National Park about one-half mile north of State Road 27, 

was particularly important in the biological exploration of the Royal Palm 

Hammock-Long Pine Key area. C. T. Simpson and companions (Simpson 1920: 

130-132) foHowed the Camp Jackson trail on what was probably the first 

significant scientific visit to Royal Palm Hammock in December 1903, as did Smal1 

in most of his early botanical work in the area (Small 1916b: 168-169). Many of 

the early naturalists knew Krome and evidently had access to his maps, for the 

depiction of "Long Key" on the maps published by W. E. Safford (1919: facing 434) 

and Simpson (1920: end papers) is obvously indebted to Krome's survey. Event ual1y , 

the Florida East Coast Railroad elected to route its Key West extension via the 

Florida Keys rather than Cape Sable; Krome'S survey records were "lost or'stolen" 

(Tebeau 1968:77) and his observations went unpublished except for a brief article in 

an engineering journal (Krome 1904), and Long Pine Key again lapsed, briefly, into 

obscurity. 

The definitive end to its isolation occurred in 1915 when the so-cal1ed Ingraham 

Highway was extended southwestward from Homestead through Royal Palm 

Hammock (Smal1 1916b, Simpson 1923; facing 269 and 332). Shortly afterwards, at 

least by the early 1920's, but probably before that, a spur road was built from the 

Ingraham Highway to the southern edge of Long Pine Key and, thence, due west for 

about seven miles. Winter vegetable farming in the area began as soon as roads 

were available. Cornwel1 and Atkins (1975:36) state that the first farming was "in 

late 1916 or early 1917," and Howel1 (1921:259) referred to farming "on the 

Everglade prairie west of the hammock" in the winter of 1917 -18. Early farming 

was largely limited to the higher and less flood-prone prairies near the Long Pine 

Key Road, particular ly the so-cal1ed finger glades. Within our mapped area, the 

total extent of land affected by farming in the pre-park era was about 250 ha. 

Much of the farming apparently was somewhat ephemeral and the area actual1y 

cultivated seems to have varied widely from year to year depending primarily on 

water levels. However, some areas of deeper soil, such as the lower ends of the 

finger glades, may have been farmed virtual1y every winter for almost 60 years. 

Easy access and, presently, a serviceable hotel in Royal Palm State Park led to a 

rapid increase in biological investigation of the area. Smal1 (1916b) notes that the 

trip to Royal Palm Hammock, formerly about a four-day expedition from Homestead culminating in a three-mile slog across Taylor Slough from Camp Jackson, 

could now be made by automobile in an afternoon. Small continued to visit almost 

annually, though his attention was more directed to new areas opened by extension 
7 

of the Ingraham Highway toward Cape Sable, and several ornithologists (Howell 

1921, Holt and Sutton 1926) and other specialists worked in the area. However, 

additions to geographical knowledge of Long Pine Key in the 1920's and 1930's 

came mainly from explorations by the collectors of tree snails (Liguus fasciatus). 

Liguus collecting ("ligging," Clench 1931) combined the appeal of natural history, 

esthetics, and exploration and attracted many conchologists and amateur enthusiasts. R. H. Humes' (1965) admittedly incomplete list of the known collectors 

included about 200 names. In monographing the genus, Simpson (1929) wrote that 

the destruction of hammocks had "almost exterminated" Liguus, but the snail 

populations-proved to be more resilient than Simpson had thought. The history of 

Flor ida tree snails is replete with announcements of the extirpation and rediscovery of local populations, as well as of the discovery and exploration of wholly 

new areas of occurrence, such as Long Pine Key. The extreme variability of Liguus 

(with, ultimately, more than 50 named color forms, plus hybrids) held out hope that 

any of the scores of hammocks hidden in the pine forests of Long Pine Key might 

yield an undescribed population. Accurate maps of hammock locations and a 

system for identifying individual hammocks, thus, became highly desirable. Serious 

Liguus collecting in the Long Pine Key area began shortly after 1900 and several of 

the early collectors there, such as Simpson and C. Mosier (Humes 1965), apparently 

knew of and used Krome's manuscript map of Long Pine Key. However, Krome's 

map was never published, and by the 1920's the map had evidently been forgotten 

or was unavailable. Thus, collectors of that era laboriously built up their own maps 

by locating hammocks from the Long Pine Key Road and the numerous narrow 

prairies (finger glades) that penetrated the southern edge of the rocklands. Library 

files at the South Florida Research Center include copies of several of these maps. 

Date and author often are not clear, some of the maps are rather fanciful as 

regards hammock locations, and some evidently were prepared for sale to other 

collectors. The closest approach to a standard map of Long Pine Key Liguus 

localities is that prepared by W. E. Schevill and W. J. Clench, -based in part upon air 

photos taken for that purpose (Clench 1931:13), and eventually published by Pilsbry 

(1946:65). In addition to the variant maps, various schemes existed for identifying 

Long Pine Key hammocks by numbers and/or names. Clench began the method of 

numbering hammocks as a means of identifying locations where he had collected 

Liguus (pers. comm.). In addition to the numbering system, Clench assigned names 

to some of the hammocks in honor of fellow shell collectors, scientists, and 

naturalists (pers. comm.). As more hammocks were discovered the numbering and 

naming system was continued by other Liguus collectors; C. N. Grimshawe, 

R. Deckert, C. C. Von Paulsen, and Humes. In more recent years, efforts were 

made (particularly by Clench, F. C. Craighead, Sr., Humes, A. Jones, and E. C. 

Winte) to standardize the hammock nomenclature. The designations we use in this 

paper are those presented by Jones, Winte, and Bass (1981). 

In general, the first-cut lumbering of pine in the Dade County rocklands proceeded 

southward from the Miami area with the advance of the Florida East Coast 

Railroad. The pine forests of Long Pine Key were the last to be cut, and lumbering 

probably began there in 1936. A U. S. Forest Service survey in the winter of 

1935-36 (Eldredge 1938) showed no sawmills operating on Long Pine Key at that 

time, although a mill located about one mile east of the present eastern boundary 

of Everglades National Park may conceivably have received logs cut on Long Pine 
8 

Key. D. B. Beard (1938: 10) wrote, "There is not very much of the original stand of 

large pines left on Long Pine Key because lumbering operations on state property 

have been in progress for a year or two now." However, the 1940 series of aerial 

photographs (Soil Conservation Service) suggests that the lumbering moved at a 

slower pace than Beard indicated, because at that time only the areas east of the 

present Long Pine Key Campground had been cut. Sawmills were located on the 

north side of the Long Pine Key Road at Osteen Hammock Glade and at Twin 

Hammock Glade near the present site of the South Florida Research Center, and 

some activity continued at the latter mill until 1947, immediately before the 

establishment of Everglades National Park. Trails used in bringing logs to the two 

mills were extended into most parts of the pineland on the large eastern island of 

Long Pine Key and many of these were later used as fire-break roads enclosing the 

prescribed burning blocks developed by Everglades National Park. Photographs of 

Long Pine Key taken in the early 1950's (Robertson 1953) indicate that the pine 

forest was clear-cut, the usual lumbering practice in southern Florida, leaving only 

cull trees of the original stand. Parts of the area may have been cut-over a second 

time removing pines that were originally considered unmerchantable. Thus, 

Cornwell and Atkins (1975:37) refer to lumbering during World War II which "cut 

out everything that would make a two-by-four." It has commonly been said 

(Robertson 1955:81, et al.) that all the pine forest on Long Pine Key was lumbered, 

except the small area at the east end of the Long Pine Key road (now heavily 

invaded by hardwoods, see D. L. Taylor and A. Herndon 1981, Fig. 1 and 2) which 

was within Royal Palm State Park. However, field work for this report iI1dicated 

that lumbering did not reach several remote areas of the main part of Long Pine 

Key (north edges of prescribed burning blocks C and D) nor any of the outlying pine 

islands west of the west end of the Long Pine Key Road. We could find no cut 

stumps nor evidence of former logging trails in these areas. We estimate the total 

uncut area at approximately 1667 ha. In some of the stands concerned, such as 

west of Deer Hammock, the mature pines were probably too small to be worth 

cutting. Other areas, such as east of Sisal Pond, now have even-aged, younger 

stands which may represent relatively recent pine invasion of adjoining rocky 

glades, perhaps related to a decrease in average water levels. However, the uncut 

area includes fairly extensive stands of mature appearance with pines of 25-35 cm 

dbh which the timber cutters must have considered were too difficult to reach. 

Until the early 1950's, farming in the Long Pine Key area was limited to prairies 

and (contra Cornwell and Atkins 1975: front is.) did not involve rocklands to any 

extent. Pr incipally because of the long-established winter agriculture, a large 

area, the so-called Hole-in-the-Donut, was left outside of Everglades National Park 

as originally established. Beginning in about 1954, application of newly-developed 

technology for rockland farming led to a rapid, massive extension of the agricultural area into virtually all of the available uplands of Long Pine Key. Thus, 

between 1954 and 1969 about 2000 ha of rockland were farmed. Preparation of the 

rockland sites for agriculture -involved removal of the natural vegetation (mostly 

thin-soil prairie and cut-over pineland, but including several small hammocks) and 

pulverizing the limestone substrate to a depth of several feet. As C. E. Hilsenbeck 

(1976) and R. E. Meador, II, (1977) have discussed, these alterations profoundly 

affected plant growth potential of the sites, once they were no longer under 

agriculture. After an extended controversy, the Hole-in-the-Donut was purchased 

by the United States and added to Everglades National Park. The last farming in 
9 

the area occurred in the spring of 1975. Management of the former agricultural 

lands has continued to be a significant resources problem for Everglades National 

Park. 

The latest historical event which has exerted a large-scale effect upon the 

vegetation of Long Pine Key was the initiation in April 1958 of a prescribed 

burning program by Everglades National Park. This activity, undertaken as a result 

of the work of W. B. Robertson, Jr., (1953) and others, has led to repeated, 

deliberate burning of ten largely pine-forested blocks comprising the major part of 

Long Pine Key. The rationale, history, and (in part) gross effects of the prescribed 

burning were reviewed recently by Taylor and Herndon (1981) and need not be 

considered further here. However, it should be noted that the prescribed burning 

program has applied principally to the parts of Long Pine Key that were enclosed 

by logging trails and more recent roads. Only accidental or incendiary man-caused 

fires and lightning fires have affected the outlying westerly pine islands. 

The name Long Pine Key seems to have replaced the ear lier Long Key during the 

1920's, although both terms continued in use for some time. The earliest reference 

to "Long Pine Key" that we found was by A. H. Howell (1921: 252 et seq.). Simpson 

called the area Long Key in 1920 and Long Pine Key in 1923 and later (Simpson 

1920, 1923, 1929). The change may have been dictated by the existence of a 

number of other places in Florida called Long Key, such as the island of that name 

in the central Florida Keys. However, the gain in precision was marginal, because 

Long Pine Key has rather frequently been confused with Big Pine Key in the lower 

Florida Keys. 

PREVIOUS WORK 

Knowledge of the flora of the southern Florida rock lands, only fragmentary prior to 

1900, developed rapidly thereafter, as over land access to areas south of Miami 

became easier. Although others contributed (see Harper 1927, Robertson 1955), 

botanical exploration of the region was primarily accomplished by Small who first 

collected there in 1901 and continued almost annual visits until the early 1930's. 

The results of his work in southern Dade County are embodied in his Flora of Miami 

(1913) and Manual of the Southeastern Flora (1933 and earlier eds.), volumes 

dealing particularly with the ferns, shrubs, and trees, an~ numerous descriptions of 

plants new to science or to the United States flora. Small briefly visited rockland 

areas west of Taylor Slough in May 1904 (Small 1904b), and, after being turned 

back by high water on an attempted trip to Long Pine Key in October 1906 (Small 

1907), he returned there on a two-week collecting trip in January 1909 (Small 

1909). The account of the trip suggests that the collecting party ranged widely 

reaching "the southwestern extremity of Long Key" (op. cit.:50) and crossing the 

main rockland area at several points. However, the coverage of Long Pine Key 

appears, quite understandably, to have been far from complete. Several plants 

conspicuous locally (e.g. Hypelate trifoliata) are not included in Flora of Miami and 

Small did not refer to such features as the Mahogany Hammocks, the royal palms 

(Roystonea regia) of Small Hammock and Little Royal Palm Hammock, nor the socalled rock reefs (linear elevations of limestone 2 to 5 m wide standing about I m 

above the surrounding marsh) which extend for miles around the western parts of 

Long Pine Key. Also, he later discussed (Small 1917) a trip to the present Osteen 
10 

Hammock, largest on Long Pine Key, as if it was his first visit to the area. Small 

later made a number of brief visits to Long Pine Key, but, as far as can be 

determined from his published reports, the 1909 venture was his only extended 

collecting trip to the area. Various later workers, particularly University of Miami 

botanists, W. M. Buswell, T. A. Alexander, and R. O. Woodbury, collected in the 

area, and in 1950-52, Robertson amassed a considerable collection primarily from 

Long Pine Key. From about 1954 until the early 1970's, CraigQead, Sr., collected 

plants very extensively in Everglades National Park, including Long Pine Key. 

Craighead's work resulted in a floristic list for the three southernmost Florida 

counties (Craighead and Lakela 1965) and contributed importantly to Flora of 

Southern Florida (Long and Lakela 1976), currently the standard manual for the 

region. In recent years, G. N. Avery and L. L. Loope did considerable collecting in 

the course of preparing their preliminary list of vascular plants of Everglades 

National Park (Avery and Loope 1980). We follow their nomenclature in this 

report. 

Not surprisingly, because he saw the entire area when it was virtually undisturbed 

by man, Small was also the first to enunciate many of the basic relationships in the 

plant ecology and plant geography of the southern Florida rocklands. For the most 

part, these observations appeared as passing remarks in the informal narratives of 

his collecting trips published in the Journal of the New York Botanical Garden. He 

recognized that the principal plant formations were pineland, hammocks, and 

Everglades prairies; that the upland flora was basically West Indian with the 

tropical element increasing rapidly as one proceeded south from Miami (Small 

1904a); that the tropical plants new to the United States flora were found mostly in 

hammocks, while the new endemic species grew mostly in pinelands (Small 1904a); 

and, that the occurrence of pine forest and hammocks were largely governed by 

recurring fire which was destructive to hammocks, but relatively innocuous in 

pinelands (Small 1911). Among early authors, E. A. Bessey (19l!) and Simpson 

(1920) also addressed the fire-mediated relationship between pinelands and 

hammocks, still a central question in rockland plant ecology, and Small returned to 

the subject frequently in his later, more formal, ecological papers (Small 1920, 

1929, 1930). Observing the enormous destruction of the increasingly frequent mancaused fires, particularly after large-scale drainage had lowered water levels, 

Small concluded that the entire upland area must have been covered by tropical 

broad-leaved forest before early Indians introduced fire into the area. Echoing 

these views, Beard (1938) wrote " ••• the Everglades Keys were once all hammock 

growth with intervening sawgrass glade lands," and F. E. Egler (1952:226) 

commented, "In short, the vegetation of south Florida during late Pleistocene preIndian times may have been a dense evergreen broad-leaved tropical jungle ••• " 

Robertson (1953, 1955) followed up on two traditions in the study of rockland 

vegetation. In the wake of the work of J. W. Harshberger (1914), R. M. Harper 

(1927), and J. H. Davis, Jr., (1943), he presented a more detailed view of the 

qualitative, descriptive ecology of southern Florida rocklands than had been 

available before, and he also considered the matter of fire and its effects upon 

rockland plant communities. In the latter context, he noted that the concentration 

of endemic plant species in fire-maintained communities argued for a long history 

\\ 

of recurrent fire, that Jightning fires appeared to be sufficiently frequent to 

provide an adequate ignition source, and that, in the absence of fire, invasion of 
11 

pinelands by hammock hardwoods proceeded rapidly enough to alter the vegetation 

of sites within periods of 10 to 15 years. Based on these points, Robertson argued 

that the then-current National Park Service management policy of excluding or 

extinguishing all fires would lead quickly to the replacement of pinelands by young 

forests of hammock hardwoods with relict pines. The result, after extended 

debate, was the initiation in April 1958 of a program of prescribed burning of Long 

Pine Key pinelands, the first such program to be undertaken by the National Park 

Service. The prescribed burning has continued with frequent up-dating of procedures, as stated most recently in the present Everglades National Park Fire 

Management Plan (1979). 

The past decade, especially since the establishment of the South Florida Research 

Center, has seen a large increase in research pertinent to rockland plant ecology 

with a move in the direction of more detailed, quantitative studies. Craighead 

(1971, 1974) addressed the familiar question of the dynamics of hammock-pineland 

relationships. Loope et al. (1979), and I. C. Olmsted, Loope, and Hilsenbeck (1980) 

presented floristic analyses of southern Florida pineland and hammock vegetation, 

both studies including stands on Long Pine Key. R. H. Hofstetter (1973) and 

Hofstetter and F. Parsons (1975) commented extensively on rockland vegetation in 

their review of fire effects in southern Florida pineland and marsh communities. 

Taylor (1981) summarized Everglades National Park fire history, and Taylor and 

Herndon (1981) analyzed the effects of long-term prescribed burning on the 

pineland shrub understory, utilizing study plots that Robertson, C. W. Senne, 

L. Chamberlaine, and others had established in the 1950's and 1960's. Hilsenbeck 

(1976) reported on the revegetation of former Long Pine Key agricultural lands 

with particular reference to invasion of these areas by Brazilian pepper (Schinus 

terebinthifolius) and Loope and V. L. Dunevitz (1981) studied the effects of Schinus 

invasion upon rockland pine forests. Additional work is in progress in several of the 

above areas. 

METHODS 

Vegetation Map 

Aerial photography, flown in February 1980 at a scale of 1:7800, was used as the 

basis for determining the distribution of plant communities. A geographically 

controlled base map, drawn from U. S. Geological Survey orthophotomaps, was 

prepared at 1:15,000 by USGS, and the communities identified from the aerial 

photography were transferred to the base map using a MAP-a-GRAPH projector. 

Field checking was done on foot and from a helicopter between February and April 

1981. 

Numbers and Names of Hammocks 

The Liguus collectors were the first naturalists to visit most of the hammocks. 

They started numbering the different hammocks as well as naming them. Over the 

years, some of the names and numbers have been changed and/or mixed up. To 

determine the exact identity (location, name, and number) of each hammock we 

referred to the original sources. The locations of the numbered hammocks on our 

map are based on the maps of W. S. Schevill, Grimshawe, M. Ross, Jr., and 
12 

Craighead, Sr., in library files of the South Florida Research Center. The 

hammock names that we used are based on those of Clench (pers. comm.) and 

Craighead {1974:56-57}. We have provided a list of all the hammocks, using the 

numbers as they appear on the map and indicating the names that go with each 

number (Appendix I). 

Elevation Survey, Soils, and Hydrologic Parameters 

A transect along which elevation was determined at intervals of 30 m was surveyed 

for a length of 2 km from Osteen Hammock to Redd Hammock (Fig. 1), traversing 

pineland, hammocks, and glades. A Dumpy level was used as the surveying 

instrument. The elevations were sighted to benchmarks on culverts along State 

Road 27 (map). 

Between 1979 and 1981, at irregular time intervals, groundwater level was 

measured with a meterstick in two wells, near Osteen Hammock and near Redd 

Hammock. These wells are located close to roads for easy access in case of fire in 

one of the hammocks. Soil depths were measured along the transect at 30 m 

intervals. 

Quantitative Vegetation Analysis along Elevation Transect 

Along the ilevation transect, vegetation plots were placed every 60 m. Ten subplots, 1 m in size, were sampled at each of 36 locations. Species' cover was 

visually estimated in each ~uare meter, and frequency of all species determined. 

In circular plots of 100 m , all the pines were counted, dbh taken, and height 

measured with a clinometer. Within the same circular area all species were 

identified. The plot numbers are indicated on the elevation transect (Fig. 1). 

Plants were classified as woody, forb, and graminoid. Woody vines and suffrutescent plants such as Vitis rotundifolia, Passiflora suberosa, and Morinda royoc were 

considered forbs. Polar ordination (Bray and Curtis 1957) was employed to group 

the vegetation along the transect. 

Hammock Fire History 

Fire history of hammocks was determined as accurately as possible from stereoscopic viewing of aerial photography. The air-photo series reviewed were black 

and white photography for 1940 (1:40,000), 1952 (1:20,000), 1960 (scale not known), 

1964 (1:30,000), and color photography (1:7800) for 1980. In some years, especially 

1960 and 1964, some hammocks were covered by clouds or otherwise obscured in 

the photography so that coverage was not complete. It should be noted that this 

method allows detection only of effects of major fires which kill a substantial 

portion of the canopy. Ground fires which did not kill much of the canopy cannot 

be detected. Similar methods were used by Loope and N. H. Urban (1980). The fire 

impact code used was as follows: M = Hammock has mature appearance on aerial 

photograph; PM = Hammock has mature appearance on aerial photograph, except 

for small shrubby portions; 1M = Hammock appears to be in condition approaching 

maturity on aerial photograph; R = Hammock exhibits signs of progressive recovery 

from burn in previous intervals; D = Destroyed; B = Evidence of burn during 
13 

interval. The difference between 1M and R is age of recovery, with R being in an 

earlier state. Change of size or shape of the hammocks was also considered in the 

analysis. 

Distribution of Shrub Stratum Plants in Long Pine Key Pinelands 

Information on the shrub flora of Long Pine Key was drawn from published lists 

(Robertson 1953, 1955; Loope et ale 1979; Taylor and Herndon 1981), from the 

vegetation transect mentioned above and other field work undertaken specifically 

for this study, and, particularly, from plant lists for 73 one-tenth acre (c. 0.04 ha) 

plots established in 1958-1965 to study the effects of prescribed burning on 

pineland vegetation. These permanently marked plots (for locations see Taylor and 

Herndon 1981, Fig. 3) measuring 33x132 feet (c. 10x40 m, not "12x48 mil, Taylor 

and Herndon 1981:5) were placed at intervals of about one-half mile along the 

interior fire roads of the main part of Long Pine Key. For convenience we have 

used the shrub lists compiled by Robertson when the study plots were originally 

established. Taylor and Herndon (1981) found that prescribed fires and other 

environmental events since 1958 have had relatively little effect upon the \ ( 

composition and diversity of the pineland shrub understory of Long Pine Key. 

In general, we have followed the categories established in an earlier report on the 

flora of southeastern Florida pine rocklands (Loope et ale 1979) in defining shrublayer plants as the native woody plants and palms occurring beneath an overstory 

of pines and reaching, at least on occasion, an erect stem height of .5 m or more on 

such sites. We have tried also to consider only areas of ecologically functional 

pineland, thus omitting stands consisting of relict pines in continuous-canopy young 

hardwood forest. Understory vegetation at such sites may often include species 

not found in more exposed areas of pine forest. Under our general definition we 

exclude: about 10 non-native shrubs and small trees of fair-ly frequent occurrence, 

of which only Schinus is abundant and widely distributed in pinelands; some 15 

woody vines and sprawling shrubs (e.g. Chiococca sp., Morinda royoc, Smilax sp.); 

about 6 low-growing woody plants (e.g. Crosso etalum ilicifolium, Licania 

michauxii); and approximately 20 suffrutescent species e.g. Eupatorium odoratum, 

Ludwigia sp.). Lantana camara, Leucaena leucocephala, and Waltheria americana, 

minor components of the pine forest shrub understory in a few areas, are also 

excluded here as being of ruderal habit often indicative of site disturbance. We 

depart from the treatment of Loope et ale (1979, Table 1) in considering that 

Eupatorium villosum is a shrub by the above criteria and that Lantana depressa and 

the regional pineland species of Hypericum are not. 

RESULTS 

Vegetation Map 

The actual area of Long Pine Key extends beyond the western border of the map to 

the vicinity of the Mahogany Hammocks ( 10 km). The present map covers an area 

of 15 x 7 km. The pineland is more or less bounded on three sides by paved roads; 

State Road 27 on the north side, the Long Pine Key Road on the south side, and a 

connecting road on the east side. A system of logging roads was built in the 1930's 

and 1940's and many of these have been maintained by the National Park Service as 
14 

fire roads. Other fire roads were added by the park in the late 1940's and early 

1950's, particularly on the west side of Long Pine Key. Roads that are presently in 

use are shown on the map. Some of the old logging roads were not maintained and 

are now grown over. The pineland proper is surrounded by prairie on most sides and 

to the south by the "Hole-in-the-Donut," an area of abandoned farmland. 

The Long Pine Key campground is located on the eastern half of the map and is 

used extensively during the dry season. 

Pinelands 

The major pineland areas of the map consist of open stands of slash pine (Pinus 

elliottii, var. densa). Most of the pine stands are more or less even-aged, having 

become established since the logging operations were ended in the late 1940's. 

The understory is typically very rich in hardwoods, most of which are tropical. 

Typical temperate woody species are Rhus copallina, Myrica cerifera, Hex cassine, 

Persea borbonia, and the palm Serenoa repens. Some tropical hardwoods are 

Dodonea viscosa, Guettarda elliptica, Psidium longipes, Bumelia salicifolia, and 

Myrsine floridana. The shrubs are mostly 1-3 m tall under the current prescribed 

fire regime. In the prescribed burn areas, the pineland has an "open" look to it: 

the pines are 12-18 m tall and are fairly widely spaced, with a low shrub 

understory. 

The ground layer consists of many herbaceous and suffrutescent perennials which 

grow in minute amounts of soil and are perpetuated by fire. It has special interest 

because it contains a number of endemic taxa including Dyschoriste oblongifolia 

var. angusta, Phyllanthus pentaphYllus var. floridanus, Tra,ia saxicola, Chamaesyce 

porter iana var. porter iana, and Poinsettia pinetorum Loope and Avery 1979; 

Loope et al. 1979). Several of the prominent herbs in this layer are graminoids, 

among them Schizachyrium rhizomatum and Andropogon cabanisii. Grasses usually 

grow in areas with a little more soil accumulation than the almost bare rocks forbs 

grow on. 

The pineland of the Miami Rock Ridge is floristically the most diverse plant 

community of Everglades National Park. The microtopography of the limestone 

with solution holes and pinnacles under the prevailing climate provide more niches 

and a habitat for about 200 species. Proximity of tropical hardwood hammocks and 

the transverse glades help increase this number by several more species at 

ecotones. Affinities of the pineland species were described by Loope et ale (1979). 

The diversity index for different areas within the pineland shows that the western 

half of the pineland is more diverse than the eastern half (Taylor and Herndon 

1981). This aspect will be discussed separately under "Distribution of Shrub 

Stratum Plants in Long Pine Key Pinelands". 

The western edge of the mapped pineland area is at a slightly lower elevation than 

most of the pineland and has a slightly different pattern of understory. While the 

species are still similar, larger areas with only graminoid cover occur here. The 

understory shrubs are not scattered evenly everywhere, but tend to grow in clumps 

on local elevations of limestone. 
15 

The ecotones between pineland and transverse glades are not always sharp. In 

several areas scattered pines extend into the prairie. Sometimes large areas of 

prairie are enclosed by pineland that is thinly stocked with pine towards the prairie 

side. Lowering of the water table during the past half century has probably made 

the ecotone between the two communities wider in some places. 

Pineland with Tall Hardwood Understory 

Prescribed burns and lightning fires do not always burn areas completely. SmaU 

corridors of pineland between two hammocks (as between Osteen and Walton, map) 

or between a road and a hammock (as Palma Vista 1 and the Long Pine Key Road, 

or Dark Hammock and an adjacent logging (fire) road) may escape fires and the 

hardwood understory grows up to hammock height. Instead of 1-3 m, the 

understory is 6-12 m tall, a deep layer of pine duff covers the ground, and forbs 

and endemic shrubs are absent (Loope and Dunevitz 1981). In such areas, pine 

reproduction is inhibited, the relict pines become senescent; and, in time the areas 

could become hammocks if fire is kept out. However, such places are few on Long 

Pine Key. 

Tropical Hardwood Hammocks 

The mapped area shows 120 numbered or named hammocks and many others that 

were never named. These hammocks are mixed stands of predominantly tropicaJ 

hardwoods that form a closed canopy at from 5-12 m, with some trees extending to 

18 m. The shrub understory is made up of smaU trees and bushes reaching to 5 m. 

The groundcover is very sparse with several graminoid species and some herbs. 

Hammocks and the pineland shrub understory have a number of tree species in 

common. 

Dominant tropical hardwoods are Lysiloma latisiliguum, Bumelia salicifolia, 

Nectandra coriacea, Exothea paniculata, Coccoloba diversifolia, Prunus myrtifolia, 

and Simarouba glauca. Olmsted, Loope, and Hilsenbeck (198 1) found several 

hammocks to be in different successional stages sin2e disturbance by fire and 

hurricane. They found a total of 42-55 species/300 m , of which 18-22 were tree 

species. 

Hammocks have a more or less continuous layer of organic soil (mostly duff) that 

averages 12-15 cm deep. The closed canopy vegetation and the moist soil layer 

create a more humid microclimate in the hammocks than in the adjacent pinelands. 

Because of this, the hammock interiors usuaUy do not burn, except under extremely 

dry conditions, but fires burning around the hammocks prune edge plants along the 

interface with adjoining pineland or prairie. 

Hardwood Scrub Without Pine Overstory 

At the edges of the pineland, in areas where glades are adjacent to pines, a sparse 

to dense hardwood scrub forms which differs somewhat from the hardwood scrub 

under pine. The substrate in these areas is often more perforated with solution 

holes than in the pineland proper. Together with the moisture difference this 

substrate change encourages hardwood scrub of certain species rather than pines. 
16 

Metopium toxiferum, Conocarpus erectus, Myrica cerifera, Hex cassine, and Persea 

borbonia are the most common species in this vegetation. The scrub does not 

usually grow taller than 6 Xn, but is mostly shorter. 

Another type of hardwood scrub occurs in early succession after fire in tropical 

hardwood hammocks. In addition to the species mentioned before, we find Trema 

micranthum, Quercus virginiana, and Bumelia salicifolia. This successional growth 

can consist of trees and shrubs. up to 6 m tall or may be of much lower scrubby 

stature. In most cases, a portion or all of the hammock floor is covered with the 

weedy bracken fern, Pteridium aguilinum. This species has allelopathic characteristics (Gliessman 1976) and can occupy a site for a number of years and exclude 

other species from establishing. 

Hardwoods, Successional in Disturbed Areas 

In some of the southern portions of the finger glades, just north of the Hole-in-theDonut, the marl prairie was cultivated for many years and then abandoned in 1947 

with the estabalishment of Everglades National Park. The hardwood vegetation 

that has become established in these areas is composed of some typical bay head 

species such as Persea borbonia, Myrica cerifera, Magnolia virginiana, Hex cassine, 

and the exotics Schinus terebinthifolius and Psidium guajava. Since these areas are 

lower in elevation than the pineland, other hardwoods rarely invade. This hardwood 

association has a very short average hydroperiod of up to one month. 

Some successional hardwood associations in the Hole-in-the-Donut are similar to 

these former glades. 

Muhlenbergia Prairies 

Composition and species abundances vary from site to site. This extensive prairie 

type occupies extensive tracts south, west, and north of Long Pine Key and the 

finger glades within the pineland. 

Muhlenbergia filipes is the dominant grass, with Cladium jamaicense (sawgrass) as 

a constant associate. About 20 other grass or sedge species occur in this 

community, as well as 60-80 forb species. Even though the species of forbs are 

more numerous than the graminoids, the grasses and sedges are by far the dominant 

plants in' cover and biomass. The mar I substrate is generally thin between 5 and 

20 cm, though there are some solution holes with deeper soils. As compared to the 

Muhlenbergia prairie described and analyzed in Taylor Slough (Olmsted, Loope, and 

Rintz 1980), the prairie of the Long Pine Key area is more variable in composition 

and cover. The finger glades are floristically more diverse than the Taylor Slough 

prairie. There are many solution holes which contain pure sawgrass vegetation. 

In places with slightly lower elevation, sawgrass is dominant because the hydro!. 

period is longer. The average hydroperiod for Muhlenbergia prairie is 2-4 months 

in northern Taylor Slough, but is less in the finger glades. Analysis of the 

Muhlenbergia prairie in the finger glades will be discussed in a later section. Some 

smalJ vegetation stands of pure tall sawgrass around willowheads and stands of 

Phragmites australis in depressions in the Muhlenbergia prairie have not been 

separated as different communities because of their infrequent occurrence. 
17 

Prairie with Scattered Scrub (Hardwoods) 

The prairie portion of this community is the same as described under Muhlenbergia 

prairie, but in a few areas where marl is either discontinuous or pinnacle rock crops 

out and the surface is slightly elevated, some hardwoods have invaded the prairie. 

These are usually short specimens of Persea, Metopium, Myrica, Conocarpus, and 

Hex. Some of these places may only have become invaded since the lowering of the 

water table during this century. There are no data indicating such lowering, but it 

is assumed that the drainage of south Florida has had an effect on the pineland as 

well. 

Prairie Species on Former Agricultural Land 

Some of the finger glades that were only slightly disturbed by farming (i.e. had 

surface rows and furrows plowed, but were not rock-plowed) still show those rows 

today. The prairie vegetation is still mostly intact in such areas. 

Cypress Prair ie 

The cypress prairie adjoins Muhlenbergia praIrie on the west side of the mapped 

area. It is essentially a transition between a Muhlenbergia prairie and a sparse 

sawgrass marsh with stunted cypress trees (Taxodium ascendens). Sawgrass is more 

often dominant in this community, mostly because the average hydroperiod is 

longer (up to 6-7 months). The density of cypress densities varies considerably and 

is lower than that found for the same community in the Big Cypress National 

Preserve (Gunderson and Loope 1981). The marl soil is slightly deeper than in the 

Muhlenbergia prairie. Cypress is at its southern limit in this area, and its stunted 

growth can be attr ibuted to the less than optimal edaphic conditions for this 

species. 

Cypress Forest 

Whether there are two distinct species of cypress in South Florida is not clear, but 

the morphologically different forms known as bald cypress and pond cypress both 

occur in the mapped area. Taxodium distichum (lib aid cypress") grows very tall 

(20-35 m) in the Fahkahatchee Strand and similar swamps. T. ascendens or 

1. distichum var. nutans (llpond cypressll) is smaller and often stunted. Abundant 

throughout the Big Cypress Swamp, it has a limited distribution in Everglades 

National Park. "Cypress forest," as designated on the accompanying vegetation 

map, includes diverse cypress vegetation which can be grouped as domes and heads. 

A dome is typically circular and has a tree height distribution that appears domelike, with the shortest trees on the margins and the highest toward the center. 

Domes occupy bedrock depressions. The elevation of the soil surface is normally 

lower in the center of domes. The substrate is peat or muck and sometimes peaty 

mar Is (Hilsenbeck et al. 1979). 

Another type of cypress community will be tentatively referred to here as "cypress 

head." This term is admittedly less than ideal since Davis (1943) used the term in a 

completely different sense--to refer to a dome connecting to a strand. Neverthe-
18 

less, we use the term to describe cypress communities which are circular in shape, 

but with a higher soil surface at the center than at the margins. Hydroperiods may 

be very short in these heads--perhaps no more than 1-2 months. 

Willowheads 

Willowheads are very scarce in the mapped area. They may be round or elongate in 

shape, have peat of 1-2 m depth, often have a deep water hole or pond in the 

center, and are near ly pure stands of willow. Thalia geniculata (alligator flag) and 

Phragmites australis are constant associates of the willow which also supports 

several vines such as Sarcostemma clausum, Mikania scandens, and Ipomoea 

sagittata. Because of the long hydroperiod (4-8 months), more aquatic plants grow 

in willowheads than, for instance, in bayheads. 

Fire is often responsible for willowhead changes. Willows invade deep burned sites. 

They are intolerant of shade and, therefore, mostly successional or maintained by 

fire (Robertson 1955, Craighead 1971, Alexander and Crook 1973). 

Open Water 

These bodies of water are "borrow pits," created during road construction, when 

the limestone was used as fill. 

Ho le-in-the-Donu t 

The Hole-in-the-Donut is an area of land which is currently in different stages of 

succession following agricultural use. Originally pineland, prairie, marsh, and 

hammock, these areas were farmed between the 1930's and 1970's and then 

abandoned. The present weedy plant cover consists mostly of the exotics Schinus 

terebinthifolius and Psidium guajava, and an association of woody vegetation, often 

with Bacchar is sp. dominant. 

Numbers and names of Hammocks 

In Appendix I, we give the numbers and names of 120 hammocks found on Long Pine 

Key. However, many of the hammocks, especially the smaller ones, have never 

been numbered or named, and 11 of the hammocks listed in Appendix I are outside 

the boundaries of the vegetation map. Those hammocks not found on the map are 

indicated by an asterisk. 

The locations of hammocks 1-58 and 59-115 on the vegetation map are based on 

maps by Schevill and Clench (Pilsbry 1946:65) and Craighead (1974:60), respectively. Hammocks 116-120 were named and numbered by Bass. A few discrepancies in the hammock location and nomenclature were found between the two source 

maps. The most important differences involved the location of hammock 

number 34 (Turkey) and the name of hammock number 28. We have followed 

Clench (pers. comm.) and have given the hammock names and numbers used by 

Craighead in parenthesis (Appendix 1). 
19 

Elevation Survey, Soils, and Hydrologic Parameters 

The transect surveyed from the pineland just east of Osteen Hammock (/I23, map) 

to Redd Hammock (II 38a), traverses several finger glades (Fig. 1). The average 

elevation of the pineland along the transect is about 2 msl (mean sea level), but the 

portion that includes Osteen Hammock and the surrounding pineland increases to 

4 m, this is probably the highest measured elevation in Everglades National Park. 

The survey shows that hammocks can either be slightly lower, slightly higher, or at 

the same elevation as the surrounding pineland. These findings should dar ify 

questions that have existed regarding elevation differences between hammock and 

pineland sites (Craighead 1971). The finger glades are always lower than the 

adjoining pineland and, under prevailing hydrologic conditions, are the only areas 

that have a short hydroperiod during the rainy season (about 1 month). As 

indicated in Figure 1, the pineland has a very discontinuous shallow soil cover, 

while the organic soil in the hammocks has an average depth of 12-15 cm. The 

marl soil in the finger glades has an average depth of 6 cm. Water level measured 

In wells at the east and west end of the transect in 1979 and 1981 (Table 1) should 

give a good indication of general groundwater levels in the pine lands. The Osteen 

well (Table 1) showed a groundwater level 100-200 cm below the surface during the 

dry season. These low groundwater levels also suggest that the woody species are 

adapted to xeric conditions. Even during the summer of 1979, this well fluctuated 

between 100 and 132 cm below the surface. At Redd Hammock, which is 

considerably less elevated than Osteen, groundwater varied from 30 to 60 cm below 

the rock surface during the same period. Hurricane Dennis produced 17 inches of 

rain in two days in August 1981 added to already high summer water levels. This 

rainfall event brought groundwater in the Osteen well up to 60 cm below the 

surface (highest level measured between 1979 and 1981), and groundwater in the 

Redd Hammock well was up to the surface during this time. The same heavy rain 

flooded extensive low-lying pinelands farther west on Long Pine Key as deeply as 

15 cm for periods of up to 6 weeks. Some of the area flooded had been burned by a 

prescribed fire about 2 weeks prior to flooding and the fire-adapted woody 

understory species had begun to resprout. Flooding delayed recovery of the shrub 

understory in such areas by up to 6 months. The inhibition of sprouting by high 

water levels suggested that the woody understory species are adapted to more 

xeric conditions and may not have been as plentiful in low pine lands prior to 

drainage as they are at present. 

Quantitative Vegetation Analysis along Elevation Transect 

For the characterization of vegetation in pineland, hammock, and transverse glade, 

a total of 36 plots was analyzed along the transect. The majority of these samples 

were located in the pineland at different elevations. These samples show 

characteristics of the eastern portion of the pineland. 

A total of 208 species was encountered in the samples, of which 167 could be 

accounted for in the pineland samples, 114 in the glades, and 51 in the hammocks 

(Table 2{a». The greatest number of species in the hammock were woody (30). 

Though the pineland also has a large number of woody species (49), the forbs 

outnumbered all plant forms with a total of 90. Forbs amounted to 20 species in 

the hammock and 68 in the prairie-glade. Of the total number of species in the 
20 

glade the forbs ranked first with 68 and there were 20 woody species. In this 

analysis, the pineland also had two more graminoid species (total 28) than the 

glade, while only one grass species occurred in the hammock. 

A large number of species occurred in more than one habitat; pineland and glade 

shared 93 species (Table 2a). The similarity between hammock and pineland, based 

on 29 species in common, is very slight, and between hammock and glade, with 9 

species in common, even less. The dissimilarity between hammock and glade is 

easily understood. The woody species in common are occasional occurrences in 

both habitats. Since the greatest number of shared species between pineland and 

glade are in the forbs, this is also the plant form where the greatest similarity 

between the two resides. This number is particularly high for the pineland, because 

the prairie-glade forb flora is present in the low elevation pineland. The pineland 

has in structure alone more niches than either. hammock or glade. The diverse 

groundcover that thrives in the open pineland and in the open glade does not exist 

in the shaded hammocks. 

Table 2(b) shows average cover percentages for woody species, forbs, and graminoids. Because of the method chosen to analyze only shrubs and not the tall canopy 

trees in the hammocks, the major cover percentage in the canopy is not included in 

the hammock cover. One can probably assume a canopy cover of 80-100 percent. 

Following that suggestion, the total cover in the hammock would be canopy cover 

plus the woody vegetation cover in Table 2(b). The hammock vegetation is. mostly 

woody. 

The total plant cover (32%) in the pineland is also made up mostly of woody 

vegetation, but with a substantial portion in forbs and graminoids. Graminoid 

cover increases towards the lower elevation in the pineland. As might be expected, 

the vegetation cover in the glades is the lowest with an average 15 percent, of 

which 75 percent cOVer is graminoid and the remainder about equally divided 

between woody species and forbs. The woody portion in the glades vegetation can 

probably be attributed to the proximity of the pineland and may have increased due 

to the lowering of the water table during this century. Overall, the cover 

percentages in pineland and glade are low. The fairly poor nutrient levels, the lack 

of soil, the porosity of the limestone, and the seasonality of rainfall probably 

account for the lack of denser cover or more biomass in this plant association. 

The current more or less even-aged pine distribution in Long Pine Key is essentially 

the result of logging that occurred in the 1930's and 1940's. Site quality (elevation, 

substrate, water, and nutrient availability) and logging history have created a pi~ 

cover of variable stature. Table 2(c) shows the average number of pines/IOO m , 

average dbh and height as well as basal area for 19 pineland plots. The most stable 

parameter of those measured was dbh (14-15 em), while height varied from 11 to 

15 m, the lower height~being at the lower elevations. The number of pines varied 

from 2.5 to 6.6/100 m , the higher numbers occurring at the higher elevations. 

These basal area and density data are similar to those of Vernick and Taylor (1981) 

for other areas of Long Pine Key. 

Table 3 shows the cover percentages and frequencies of individual species in each 

vegetation type. Species were selected for either commonness or importance. 

Species that occurred only in one vegetation type and also had less than .1 percent 
22 

Table 4 lists the major tree and graminoid species in the pineland plots that may be 

indicative of certain environmental parameters. The occurrence of trees like 

Conocarpus, Persea, and Byrsonima in some plots and their absence in others 

suggest elevation and hydrology as a factor for this distribution. They do not occur 

in plots 1-4, but are more abundant in lower elevation plots. Lysiloma only occurs 

in one plot very close to Redd Hammock (1169). Guettarda scabra, Dodonaea 

viscosa, Metopium toxiferum, and Bumelia salicifolia are more or less evenly 

distributed in high and low pineland, while Randia aculeata and Myrica cerifera are 

more abundant in lower pineland. Plots 1-4 (the highest elevation) show 

Andropogon cabanisii, a pineland grass, but no Muhlenbergia filipes, a prairie grass. 

However, the prairie grass is abundant in low elevation pineland plots 5, 8, 9, 10, 

and 11, while Andropogon cabanisii is less com mon in these latter samples. 

Hammock Fire History 

Table 5 lists the hammocks named on the map with an indication of the fire 

damage that affected each hammock over the time period 1940-1980. Where no 

fire impact code is given for a hammock in the year of a particular air-photo 

series, either there was no aerial photography for that portion of pineland, or the 

proper status of the hammock could not be recognized on the photography due to 

cloud cover. Exact interpretation of the fire-damage status of a hammock from 

aerial photography was often difficult. Hammocks that had recently burned (B) 

were readily distinguishable in most cases, but study of park fire records and the 

entire series of available photographs was sometimes needed to distinguish 

immature (1M), recovering (R), and mature (M) stages. In the case of the 1980 

aerial photography, we were able to verify interpretations of the photographs by a 

ground check of the condition of hammocks. 

The size and shape of most hammocks seem to have stayed remarkably constant 

over the period studied. Most shifts were minor, either due to incomplete burning 

or lack of burning over an interval. In several places indicated on the map, stands 

of tall hardwoods have developed either between two narrowly separated 

hammocks or in unburned areas. 

From the list of fire impacts (Table 5) note that some hammocks (about 10) did not 

burn during the time interval considered. However, the majority of hammocks 

(about 90%) burned repeatedly, particularly between 1945 and 1960. Maxwell 

Hammock (1186) burned in 1940 and has been recovering over the last 40 years and 

is still immature. More often, we find that hammocks burned in the 1940's were 

mature in 1980. Some examples of such recovery are Osteen (1123), Gun (1124), 

Turkey (1134), Clapp (1137), and Decamp (1147). As previously discussed (Olmsted, 

Loope, and Hilsenbeck, 1981), the time required for recovery after fire and the 

composition of the recovering stand tend to vary for each hammock. 

Distribution of Shrub-stratum Plants in Long Pine Key Pinelands 

The pine forests of Long Pine Key are characterized by the presence of a diverse 

understory of shrubby hardwoods and palms beneath the pines. For example, 45 

species occurred in the pineland shrub stratum along the 2 km transect on eastern 
23 

Long Pine Key discussed above (see Appendix 11). Overall, we have recorded 61 

species that meet the stated criteria for shrubs (Table 6), and, although the list is 

based on extensive surveys, it is likely that a few additional shrubs will be found to 

occur rarely in the Long Pine Key pinelands. Species that are primarily West 

Indian in distribution comprise 75 percent (46 of 61) of the known shrub flora. Most 

of the species of more temperate distribution (10 of 15) occur typically at wetter 

pineland sites, such as in deep sinkholes or in the ecotonal belts between pine 

forest and Muhlenbergia prairie. This distribution is in line with Small's (1907) 

early observation that the upland flora of southern Florida was much more tropical 

in composition than the wetland flora. 

The 73 plots established for study of the effects of fire on the Long Pine Key shrub 

stratum (see METHODS) provide the best available measure of the evenness of 

distribution of the shrub species. Fifty-seven of the 61 species known to occut in 

the Long Pine Key pine lands at large have been found on these one-tenth acre 

(0.04 ha) plots. The missing species are either extremely rare in the pineland 

understory (Crossopetalum rhacoma, Diospyros virginiana, Simaruba glauca; each 

recorded only one or two times), or (Alvaradoa amorphoides) are more common but 

very locally distributed on Long Pine Key. 

Constancy of occurrence for each species found on the 73 study plots is shown in 

Table 6. In all, 21 species occurred on more than half the plots, 7 species on 26 to 

50 percent of the plots, and 29 species (51 % of the total) on less than one quarter 

of the plots. The 28 most constant shrubs included 21 West Indian and 7 more 

temperate species: the 29 least constant included 22 West Indian and 7 more 

temperate species. Of the least constant group, only a few species (notably 

Bourreria cassinifolia, Colubrina cubensis, and Hypelate trifoliata) are also rare in 

the flora of Long Pine Key at large. The remainder either have patchy, local 

distributions in the pineland shrub stratum, or they are typically plants of tropical 

hammock or wet-site thicket vegetation that occur only sparingly in pineland. 

Number of shrub species per 0.1 acre plot was continuously distributed from 14 

through 29 with a mean diversity of 22.1 species. Twenty-two plots (30%) 

had 14 through 19 species of shrubs, 32 plots (44%) had 20 through 24 

species, and 19 plots (26%) had 25 through 29 species. The principal trends 

evident in the plot diversity data were tendencies toward decreasing diversity from 

upland to less elevated sites, and, within the upland sites, toward increasing 

diversity westward in the portion of Long Pine Key sampled by the plots. Most of 

the plots that had fewer than 20 shrub species were located in the transition belts 

between pineland and Muhlenbergia prairies. At the upper end of the diversity 

scale, 12 of 27 plots in A and B, the westernmost prescribed burning blocks (see 

Taylor and Herndon 1981, Figure 3), supported 25 or more species of shrubs as 

opposed to 7 of 46 plots in the more easterly prescribed burning blocks. 

The somewhat greater diversity of the shrub stratum in western parts of Long Pine 

Key is associated with fairly obvious, but unexplained, differences in the limestone 

substrate. Over the eastern two-thirds of the major upland area of Long Pine Key, 

the surface of the Miami Oolite is extremely irregular with angular "dog-tooth" 

ridges and projections enclosing shallow solution cavities. By contrast, sizable 

areas in blocks A and B are characterized by monolithic exposures with relatively 
24 

smooth surfaces and numerous large, well-like solution holes up to 3 m deep. In 

these characteristics the surface limestone is more similar to that of the Lower 

Florida Keys than it is to exposures in other pine rocklands of the southeastern 

mainland. Presumably, such present differences may indicate local variations in 

depositional environment or subsequent solution, but no relevant studies of the 

Miami Oolite seem to be available. The nature of the limestone affects shrub 

understory composition and diversity in two ways: the deep, more or less 

permanently wet solution holes support a more general distribution of such wet-site 

species as Salix caroliniana and Annona glabra than is found elsewhere on Long 

Pine Key; and, several of the less common pineland shrubs are largely limited on 

Long Pine Key to the areas of smoother, more massive surface limestone. These 

species include Alvaradoa amorphoides, Bumelia celastrina*, Colubrina arborea*, 

Colubrina cubensis, Hypelate trifoliata*, Jacguinia keyensis*, and Mastichodendron 

foetidissumum. We can see no adequate ecological explanation for the restriction 

of these West Indian species to a particular facies of limestone on Long Pine Key, 

but it is notable that several (starred above) occur elsewhere in southern Florida 

primarily in coastal or beach ridge hammocks. This suggests that their occurrence 

on Long Pine Key may be relict, perhaps dating from a higher, presumably postglacial, sea st~nd when Long Pine Key uplands may have been in closer contact 

with coastal plant communities. The abundance of Conocarpus erectus throughout 

the pineland-Muhlenbergia prairie ecotonal belts surrounding Long Pine Key may 

also be relevant in this regard. 

Counts of the number of hardwood stems more than 3 feet high are available for 

104 areas of 0.025 acres (0.01 ha) within the 73 plots. These data show that any of 

19 species may be the most abundant hardwoods in local areas of the Long Pine 

Key pineland shrub stratum. The 11 most frequent numerical dominants and the 

number of areas in which each was the most abundant hardwood species were: 

Guettarda scabra, 30; Myrsine floridana, 15; Ardisia escallonioides, 9; Conocarpus 

erectus, 8; Bumelia reclinata, 6; Dodonaea viscosa, 5; Guapira discolor, 5; 

Byrsonima lucida, 4; Hex cassine, 4; Chrysobalanus icaco, 3; and Myrica cerifera, 3. 

The commonness of Myrica was also evident in the analysis of pineland between 

Osteen and Redd Hammock (Table 3). Most sites where Conocarpus, Bumelia, Hex, 

Chrysobalanus, or Myrica was the most numerous hardwood were located in low 

pineland transitional to Muhlenbergia prairie (see also Table 4). On higher pineland 

sites, Guettarda scabra was ubiquitous in occurrence over the eastern sections of 

Long Pine Key, where it was the most common hardwood on 38 percent of the 

areas sampled (27 of 71, see Table 3). Farther west in prescribed burning blocks 

A and B, the species is generally absent or rare and it was the most numerous 

hardwood on only 9 percent of the sample areas (3 of 33). Less marked local 

variation in the abundance of particular species is widespread on Long Pine Key, 

usually without obvious explanation. It seems likely that some of these patterns of 

occurrence may reflect local variation in fire effects summed over periods of 

decades. Taylor and Herndon (1981) detected numerous, but mostly minor, changes 

in relative abundance of shrub understory species over a 22-year period and 

interpreted these as fire-related. Thus, although most plants in the Long Pine Key 

shrub understory seem to be old individuals that have survived many fires, slight 

differences in fire tolerance among species may result in long-term fluctuations in 

understory composition. 
25 

SUMMARY 

1. We describe the history of discovery, exploration, and use of Long Pine Key 

in Everglades National Park. 

2. A vegetation map of Long Pine Key is provided. The variations of pineland, 

tropical hardwood hammock and Muhlenbergia prairie in composition, structure, and environmental parameters are discussed in detail. 

3. A 2 km elevation transect from Osteen Hammock to Redd Hammock shows 

the distribution of plant communities and soil depths along its length. 

4. We trace the names and numbers of 120 tropical hardwood hammocks, 

appearing on the map, to original sources. 

5. We give the results of quantitative analyses of 36 vegetation plots (10 x 1m2) 

along the elevation transect. Species plant cover is estimated and frequency 

calculated. 

6. A total of 208 species occurred along the transect, with 168 in the p,ineland, 

114 in the prairie glades, and 51 species in the hammocks. 

7. We discuss the peculiarities of the understory shrub distribution and relate 

their occurrence to substrate differences. 

8. The fire history of the tropical hardwood hammocks between 1940 and 1980 is 

traced from aerial photography and helicopter surveys. 

, 
26 

CHAPTER 2 

PINELAND SHRUB UNDERSTORY: COMPARISON OF 

LONG PINE KEY WITH OTHER SOUTHERN FLORIDA PINE FORESTS 

As noted above, the pine forest of Long Pine Key is characterized by the presence 

of a remarkably species-rich understory of shrubby hardwoods and palms. Here we 

compare the pineland shrub stratum of Long Pine Key with that found in other 

regional pine forests extending from the Lower Florida Keys to Big Cypress 

National Preserve. 

METHODS 

Our comments on variation in shrub species diversity between stands of pine forest 

in the southern Florida region build upon the recent study of rockland pine forest 

floristics by Loope et al. (1979). These authors compared the flora of seven sites 

on Long Pine Key, five sites in southeastern mainland pine areas outside Everglades 

National Park, and one site on Big Pine Key based ~n the plants encountered within 

a 40 x 40 m "macroplot" and 20 quadrats each 1 m at each site. We undertake to 

present complete lists of species occurring in the shrub stratum in five geographic 

areas of southern Florida pineland: 1) Lower Florida Keys, including extensive pine 

stands on Big Pine, Little Pine, No Name, Cudjoe, and Sugarloaf Keys, and smaller 

areas on several additional keys; 2) Long Pine Key; 3) Southern Biscayne pineland, 

remnant stands of pine in the rockland area east of Taylor Slough from its southern 

extremity north approximately to Silver Palm Drive (SW 232nd Street); 4) Northern 

Biscayne pineland, similar pine remnants north of the preceding and extending to 

the southern edge of the built-up area of greater Miami; and, 5) Big Cypress 

National Preserve, principally pinelands in the southern and central parts of this 

National Park Service area located west of the Everglades in Monroe and Collier 

counties. The division between areas 3 and 4 above is necessarily somewhat 

arbitrary as the original "Biscayne Pineland" (Small 1913a) was continuous across 

this region. We have elected to make the division at the point where deposits of 

siliceous sand become conspicuous on the pineland forest floor. Typically, as one 

proceeds northward in the Biscayne Pineland, scrub oaks (Quercus minima, 

Q. pumila, Q. virginiana var. geminata) first appear in the pine forest understory a~ 

such sites. Thus, the "Navy Wells" and "Camp Owaissa Bauer" stations of Loope et 

al. (1979) are placed in the southern section and their "Thompson Park", "USDA", 

and "Tamiami Pines" stations in the northern section. 

In compiling lists of shrub species for the various areas (Table 6), we have drawn 

upon all available sources of floristic data. Our listing for the Lower Florida Keys 

pinelands depended critically upon the field notes and comments of G. N. Avery 

and included information from lists presented by J. D. Dickson, III, et al. (1953), 

Robertson (1953, 1955), and Loope et al. (1979). Listings for the two sections of 

the Biscayne pineland were derived from Robertson (1953, lists 1 and 2 of 

Table 10), Loope et al. (1979), the notes and comments of Avery and A. Herndon, 

and specific surveys undertaken by Robertson. The list of pineland shrubs of Big 

Cypress National Preserve was based on a preliminary checklist of vascular plants 

of the area (Black and Black 1980), data provided by L. Gunderson and 

R. Rochefort, and field surveys by Robertson. 
1 

\ 

27 

The thoroughness of coverage of the available pine stands varied considerably 

between the four principal geographical areas. For Long Pine Key and the Lower 

Florida Keys, the lists of shrubs (Table 6) are based upon surveys that included 

most parts of these pinelands and may be considered largely complete. Coverage 

of the poorly accessible and greatly fragmented pine areas of Big Cypress National 

Preserve (Patterson and Robertson 1981, Fig. 6) was extensive, but less thorough. 

In contrast to these areas in which the overall extent of pine forest is relatively 

little changed from that present originally, the Biscayne pineland exists only as 

small, isolated stands, more than 95 percent of its original area having been 

destroyed by land development (Loope et al. 1979). Moreover, the diversity of 

understory vegetation in many of the existing stands has been greatly reduced as a 

result of invasion by the exotic tree Schinus terebinthifolius (Loope and Dunevitz 

1981). Information on the composition of the shrub stratum of the Biscayne 

pineland was necessarily taken from a few remnant stands, particularly in Dade 

County parks, that seemed to approximate natural conditions. Present lists for the 

Biscayne pineland (Table 6), thus, probably omit an unknown number of less 

common shrubs that occurred originally. However, we feel that the likely number 

of omissions is too small to affect our general comments on the variation in 

diversity of the pineland shrub stratum within south Florida. 

RESULTS 

Our final sanitized list of plants occurring in the shrub stratum of southern Florida 

pine forests (Table 6) totalled 95 taxa. Their distribution across the region is 

discussed below. 

Regional Distribution and Diversity of Shrub-stratum Plants 

Twelve species occur in the shrub stratum of pine forests throughout southern 

Florida: Baccharis halimifolia, Bumelia salicifolia, Chrysobalanus icaco, Ficus 

~, f. citrifolia, Myrica cerifera, Myrsine iloridana, Persea borbonia, Randia 

aculeata, Rhus copallina, Sabal palmetto, and Serenoa repens. These generally 

distributed pineland shrubs include both West Indian and temperate species. Most 

are relatively conspicuous in pinelands throughout southern Florida, but several 

(Bumelia salicifolia, Ficus citrifolia and Randia aculeata in northern Biscayne 

and/or Big Cypress) are rare or local in some areas. 

Two other patterns of wide distribution are represented by the shrubs found 

throughout mainland areas, but not in pinelands of the Florida Keys, and those 

found in all areas except Big Cypress. Widespread taxa missing from the shrub 

layer of Florida Keys pine forests are: Bumelia reciinata, Callicarpa americana, 

Dodonaea viscosa, Hex cassine, Quercus virginiana, and Salix caroliniana. Most 

range extensively north of our area and only Dodonaea is known to occur in the 

Lower Florida Keys (SmallI913b, 1933; Dickson etal. 1953; Avery pers. comm.), 

where it is an uncommon small tree in hardwood hammocks (= D. microcarya 

Small). Widespread taxa missing from the shrub layer of pinelands only in Big 

Cypress are: Bourreria cassinifolia, Byrsonima lucida, Cassia chapmanii, 

Coccothrinax argentata, Croton linear is, Guettarda scabra, Lantana involucrata, 

Metopium toxiferum, Psidium longipes, and Trema micranthum. All are tropical 

forms, and, except for the widely dispersed but very rare Bourreria cassinifolia, all 
28 

are relatively prominent in the pineland shrub stratum on the Lower Florida Keys 

and Long Pine Key and tend to decrease rapidly northward in the Biscayne 

pineland. Only Metopium and Trema are reported from Big Cypress (Black and 

Black 1980) and neither has been noted there in pineland. 

The shrub stratum of southern Florida pine forests is composed predominantly (67 

of 95 taxa, 7196) of taxa derived from the flora of the Bahamas and Cuba, and, 

within southern Florida, most of these forms reach or approach their northern 

range limits. Elements of a second zonal flora, that of the pine flatwoods of the 

southeastern United States coastal plain, enter the shrub stratum of regional 

pinelands in northern Biscayne and Big Cypress at sites where the limestone 

substrate is mantled by deposits of acid sand. However, the characteristic shrubby 

plants of the pine flatwoods are represented only sparingly within our area (7 taxa, 

about 8 percent of the regional shrub-layer flora). Thus, species diversity of the 

pineland shrub stratum tends to decrease sharply from south to north within 

southern Florida (Table 7), primarily because the West Indian forms that reach 

range limits far outnumber the few flatwoods species added to the shrub stratum. 

Diversity of the pineland shrub stratum also appears to be directly related to siteto-site differences in the linear extent of peripheral and internal interface between 

pine forest and other plant communities. Throughout southern Florida, the 

pineland shrub stratum typically is more diverse at sites where pine forest encloses 

or adjoins hardwood hammocks. Bursera, Exothea, and Lysiloma exemplify. hardwoods whose occurrence as pineland shrubs is largely limited to such contact belts 

which rarely exceed 100 m in width. Sites where pine forest is in contact with 

lowland vegetation usually exhibit reduced shrub-stratum diversity, but may 

include species that are not found locally at more elevated pineland sites. 

EXamples are seen in the frequent occurrence of Taxodium in the understory of 

smaller pine islands in Big Cypress and the extensive dominance of buttonwood 

(Conocarpus erectus) in the shrub layer of low pinelands on Long Pine Key 

(Robertson 1955:110-112). Limestone sinkholes of varying width and up to c. 4 m 

deep occur throughout the pinelands underlain by Miami oolite and are particularly 

frequent in the Lower Florida Keys and Long Pine Key. Many of- the sinkholes 

intersect the ground water table and may hold water through much of the usual dry 

season. Swamp hardwoods (Chrysobalanus, Hex cassine, Magnolia, Persea) are 

often found at such sites and the occurrence in pineland of Annona, Ce halanthus; 

Salix, and Sambucus is almost entirely limited to the deeper sinkholes Robertson 

1955:114,119). 

More local characteristics of the shrub understory in particular areas of pineland 

are noted below. 

Lower Flor ida Keys 

The Florida Keys pineland differs from all other regional pine forests, perhaps from 

all other pine forests, in having a more-or-Iess continuous small-tree stratum (at 

roughly 3 to 8 m) of fan palms (Coccothrinax, Thrinax) at many sites. The shrub 

stratum beneath the pine-palm overstory is notable for the number of unique taxa 

(Table 7). More than one-third of the species are represented in southern Florida 

pine forests only at sites in the Lower Florida Keys. These include six West Indian 
29 

species not known from the southern Florida mainland {Caesalpinia pauciflora, 

Catesbaea parviflora, Pisonia rotundata, Reynosia septentrionalis, Strumpfia 

maritima, and Thrinax morrissii}, of which Pisonia and Thrinax are common and 

generally distributed in the Lower Keys pine forests; and 14 species which on the 

mainland are limited primarily to coastal plant communities. Occurrence of plants 

of the latter group in the pineland shrub stratum seems attributable to the very 

extensive linear edge between pine forest and shoreside vegetation types that 

exists in the Lower Keys. At many sites, the Lower Keys pinelands have abrupt 

junctions with areas of mangrove, coastal hammock, and beach hammock, and most 

of the plants concerned enter the pineland shrub stratum only in relatively narrow 

belts along these interfaces. However, such species as Coccoloba uvifera, Eugenia 

myrtoides, Piscidia piscipula, and Pithecellobium guadalupense are more widely 

distributed as pineland shrubs. 

As might be expected, the shrub stratum of Lower Keys pine forests is most similar 

floristically to that of Long Pine Key (Table 7), the most southerly mainland pine 

area. Notably, however, a number of West Indian species found in the Long Pine 

Key pinelands are apparently missing from pine forests of the Lower Keys .. 

Examples are: Alvaradoa amorphoides, Colubrina cubensis, Dodonaea viscosa, 

Eupator ium villosum, Hex krugiana, and Tetrazygia bicolor. Some of these 

absences may result from the vagaries of overseas establishment and subsequent 

spread within southern Florida. However, some probably are due to the more xeric 

environment of the Lower Keys pinelands, rather than to failure to reach this area. 

Some plants that are abundant and generally distributed in the Long Pine Key pine 

forests are markedly restricted to moister sites in pine forests of the Lower Keys. 

Two good examples are saw palmetto (Serenoa) and bracken (Pteridium aguilinum 

var. caudatum), both ubiquitous in mainland pine areas, but confined to shallow 

saucer-like solution depressions in the Lower Keys pinelands and virtually absent 

from the general level of the forest floor. Shrubs showing a similar distribution 

include Chrysophyllum oliviforme, Eugenia axillar is, Ficus citrifolia, Guettarda 

elliptica, Guettarda scabra, and Myrsine floridana. In all cases the contrast is 

between general distribution in Long Pine Key pine forests and variously limited 

distributions related to moister sites in the Lower Keys. Other species common in 

the shrub understory of mainland pine forests (Ardisia escallonioides, Guapira 

discolor, and Lantana involucrata) tend to occur in the Lower Keys mostly within 

hammocks or at hammock edge~. Such occurrence patterns suggest that the more 

xer ic conditions in the Lower Keys pineland affect both the local and regional 

distribution of species in the pine woods understory. 

Marked differences exist in the composition and density of shrub understory 

vegetation from site-to-site in the Lower Keys pineland. In many parts of Big Pine 

Key the .shrub stand is low and sparse, and the forest floor is scantily covered with 

grasses or consists of smooth bare limestone under the pine and thatch palm-silver 

palm layers. In such areas the shrub understory vegetation is composed of 

prostrate plants of Ernodea littoralis, Morinda royoc, and Smilax havanensis; low 

mounded bushes of Byrsonima lucidum and Psidium longipes; and widely scattered 

erect shrubs, chiefly Croton linear is, Metopium toxiferum, Pisonia rotundata, and 

Pithecellobium guadalupense. Throughout the Big Pine Key forest local aggregations of shrubs form small incipient hammock clumps, apparently at sites 

enjoying greater fire protection than the general expanse of forest. These occur on 
30 

the small rocky knolls found at the bases of some of the larger pines and around 

many of the sink holes. In contrast, the main pine area of Cudjoe Key has an 

almost continuous hardwood understory 6 m or more high in places. Here also the 

forest floor is covered with a deep mat of pine straw and dead grasses. The 

pineland shrub stratum of Little Pine and No Name keys is similar, but somewhat 

less dense. 

These differences seem to be the result of the relative frequency of recent fires, 

largely man-caused. Big Pine Key pine lands are extensive, continuous, and readily 

accessible. They burn frequently, and signs of recent fire are evident throughout. 

Pine stands on other islands of the Lower Keys are smaller and more isolated 

(Little Pine Key is accessible only by boat), often consisting of relatively narrow 

ridges surrounded by mangroves and open salt flats. Signs of former fires are 

present in all areas, but many stands have not burned for many years and the more 

advanced development of the hardwood understory reflects this fire-free interval. 

The solution holes mentioned above determine the occurrence of a number of taxa 

in the Lower Keys pine forests. Two general types occur - shallow saucer-like 

depressions in sizes to 20 m in diameter, and deep "wells" occasionally as much as 

3 m in diameter, but usually much less. The former are seasonally flooded, and 

their lower parts may stay wet throughout the year. Most of the "wells" hold 

permanent water, although the level fluctuates greatly. The plants potentially 

occurring in any sink apparently depend upon moisture relations as governed by the 

depth of the depression, i~s areal extent, and the nature of the walls (relative 

steepness of the slope to the center in the case of the saucer depressions; presence 

or absence of crevices or niches in the perpendicular sides of the wells). The 

plants, of those possible, which actually occur doubtless illustrate the chances of 

establishment and survival, especially in the case of the wells, which often provide 

room for no more than a single shrub or fern clump. 

Though there is much variety in the sink-hole vegetation, especially on Big Pine 

Key, several general types may be noted. Large saucer depressions are commonly 

occupied by sawgrass (Cladium) and ringed by buttonwood (Conocarpus) trees, and a 

mixed growth of scrubby hardwoods. Occasionally such aquatics as Typha and 

Sagittaria occur in the center. Shallower saucers may have dense beds of saw 

palmetto or a ring of saw palmetto around a central area of sawgrass. The slopes 

of many of the small saucers and the entire central area of some of the shallower 

ones commonly support growths of bracken. Higher slopes of the saucers usually 

have shrubby hardwoods ranging from a few individuals to a dense continuous fringe 

of sub-hammock growth. Several of the pineland shrub understory plants limited to 

such sites have been mentioned. 

The deeper wells provide environmental niches for several species otherwise absent 

from the Lower Keys. Among these are Annona glabra, Chrysobalanus icaco, 

Per sea borbonia, and leather fern (Acrostichum). Commonly each small well 

contains plants of only one of these species, several plants or a single large 

individual, or combinations such as one pond apple tree and a clump of leather fern. 

A few of the larger wells have more varied vegetation. 
31 

Long Pine Key 

The diverse shrub layer of Long Pine Key pine forests (Table 7) results primarily 

from the area's intermediate location in relation to the principal regional climatic 

gradients. Thus, seven shrub-layer taxa of West Indian derivation occur only in 

Long Pine Key and the Lower Florida Keys and eight only in Long Pine Key and the 

Southern Biscayne pinelands (Table 6). It seems reasonable to suppose that the 

former group of plants tends to be limited northward by sensitivity to increasingly 

frequent frost and the latter group tends to be limited southwestward along the 

gradient of decreasing rainfall by sensitivity to the much more xeric environment 

of Lower Florida Keys pine forests. Of the five pineland shrubs that (within 

southern Florida) are unique to Long Pine Key, two (Cephalanthus and Magnolia) 

are widespread wet-site species of little biogeographical significance, and three 

are West Indian forms for which Long Pine Key evidently provides the only suitable 

pineland growth sites available in southern Florida. 

Extensive edge with other plant communities and the extremely broken and eroded 

limestone substrate also contribute to the diversity of the shrub-layer in the Long 

Pine Key pinelands. The pine forest encloses' scores of hardwood hammocks and is 

enclosed by Muhlenbergia prairies from which linear extensions, the so-called 

"finger glades," penetrate the main body of pineland (see vegetation map). As 

noted, many plants of the pineland shrub-layer are particularly associated with 

these ecotones. Besides an array of wetter sites in the deeper sink holes, the 

jagged, irregular limestone forest floor of much of Long Pine Key provides 

innumerable, semi-protected micro-sites favoring establishment and survival of 

shrub-stratum plants. 

Southern Biscayne 

From scanty descriptions in the literature (Harshberger 1914, Harper 1927) and 

present remnant stands, it is evident that the shrub understory quickly becomes 

much less diverse as one proceeds north across this area. The understory of stands 

at the extreme southern end of the Biscayne pineland (such as Pine Island and 

Parachute Key just east of Taylor Slough in Everglades National Park) closely 

resembles that of Long Pine Key in appearance and diversity. Only a dozen or so 

species of shrubs are missing. Most of these are rare and local on Long Pine Key 

and may well have been present originally in pine forests at the lower end of the 

Southern Biscayne area. Eight miles to the northeast in the vicinity of Homestead 

the aspect of the shrub understory in remnant patches of pine forest is dominated 

by low palms, most of the West Indian hardwoods have disappeared, and most of 

those that persist are rare. Areas of greater shrub-layer diversity tend to be 

closely restricted to the pinelands bordering hammocks (as at Camp Owaisa Bauer, 

Loope et al. 1979), and in this region, as Small (l904a:51) reported from his first 

foray into the area, " ••• the pine lands are vast and the hammocks are few and 

small." Increased diversity through local re-appearance of some West Indian 

hardwoods in the pine forest understory may also have occurred originally in the 

narrow transitional belts around deep sink holes ("banana holes", Harshberger 1914) 

and along the transverse glades and the bordering Muhlenbergia prairies to the 

east. However, such sites have been largely obliterated by development. 
32 

Northern Biscayne 

In this area, the limestone becomes increasingly mantled by sand northward, 

several shrubs characteristic of southeastern United States pine flatwoods appear 

in the understory and attenuation of the West Indian element in the pineland shrub 

stratum continues. Species of West Indian derivation comprise about half the 

overall shrub-stratum list, but most are scarce and local and low palms and scrub 

oaks dominate the aspect of the pine forest understory at most sites. 

Big Cypress 

The pineland shrub understor y of Big Cypress is the least diverse in our area and 

approaches a balance in number of species between those characteristic of 

southern sandy pine flatwoods (6) and tropical West Indian forms (9). The former 

tend to be more widely distributed and the flatwoods element also includes several 

conspicuous suffrutescent plants, such as Rubus trivialis and Satureja rigida. 

Conversely, the West Indian shrubs tend to be limited to cold-protected sites, such 

as pine islands surrounded by sloughs. Hundreds of such sites exist in Big Cypress 

and it is likely that additional West Indian plants will be added to the shrub-stratum 

list which already includes several taxa (Ardisia and Eugenia axillar is) that aren't 

recorded for the northern part of the Biscayne pineland. 

Density Characteristics of the Pineland Shrub Stratum 

Although considerable change in the density and aspect of the shrub understory 

appears to accompany the above trends of change in diversity, few comparative 

data are available. For Long Pine Key, Taylor and Herndon (1981) summarized 

counts of stems of the taller hardwoods on a number of study plots (bottom, 

Table 8), but no comparable data base exists for other southern Florida pinelands. 

Loope et al. (1979) recorded plot counts in areas representing all the regional 

pine lands except those of Big Cypress, and information on the density of shrub 

species, extracted from their data, is given in Table 8. The scanty quantitative 

record seems to support general impressions that overall density of the shrub 

stratum and the relative predominance of hardwoods in mainland pine forests tend 

to decrease sharply northward from Long Pine Key and nearby areas of Southern 

Biscayne, while relative predominance of palms increases. In northern parts of the 

or iginal Biscayne pineland (e.g., Harshberger 1914, Plate IV) and in Big Cypress 

(Patterson et al. 1980, cover, Patterson and Robertson 1981, Fig. 26), palms 

comprised most of the shrub stratum with hardwoods prominent only locally. The 

low density of the shrub stratum on Big Pine Key (Table 8) reflects the monolithic, 

plate-like character of the limestone surface at the site studied. At Lower Florida 

Keys sites with more broken substrate, density of the pineland shrub stratum 

approximates that found at upland sites on Long Pine Key. 

In contrast to the species characteristics of sandy pine flatwoods, a large 

proportion of the West Indian and swamp-forest hardwoods, (more than half the 

species listed in Table 6) are potentially capable of reaching at least small-tree 

stature on southern Florida pineland sites. For example, Taylor and Herndon (1981, 

Table 2) listed 24 species that had attained heights greater than 10 feet (C. 3 m) on 
33 

Long Pine Key study plots, even under conditions of frequently recurring fire. 

Pineland fires reduce the height of the shrub understory without having much 

effect on understory density, because most of the hardwoods resprout readiJy from 

underground parts protected in niches of the limestone. On Long Pine Key, 

recovery of hardwoods after fire is direct and rapid, and a fire-free interval as 

short as 10 to 15 years is sufficient at some sites for establishment of the a moreor-less continuous smal1-tree understory. Management to prevent excessive 

accumulation of fuel and to preserve populations of rare and endemic herbaceous 

plants in pinelands (Loope et al. 1979) must take account of the fact that many 

species of pineland "shrubs" in southern Florida are not genetically of shrub form or 

stature and wil1 quickly become trees in the absence of fire. 

DISCUSSION 

Shrub-stratum diversity in southern Florida pine forests is greatest, around 60 

species, in the Lower Florida Keys, Long Pine Key, and (probably) the southern part 

of the Biscayne pineland and decreases northward to fewer than 30 species in the 

pinelands of Big Cypress. Diversity is diminished primarily by the loss of West 

Indian hardwoods from the understory as one proceeds north in the Biscayne 

pineland. Attenuation of the West Indian element doubtless is related in part to 

increasing frequency of damaging freezes in the open pinelands, but the great 

decrease in the amount of interface between pine forest and tropical hardwood 

forest in the Biscayne pineland as compared with Long Pine Key may be an equal1y 

important influence. A few shrubs characteristic of the pine flatwoods of the 

southeastern United States enter the southern Florida pineland flora in Big Cypress 

and northern parts of the Biscayne pineland where sand deposits mantle the 

limestone. The fact that similar species of the same ericaceous genera (Lyonia, 

Vaccinium, and Befaria) are found in sandy pine lands of western Cuba (Leon and 

Alain 1974) suggests that their occurrence in southern Florida is limited by 

substrate, rather than by climate. 

Ecologically similar forests of long-needle pines, which could be regarded as a 

single vegetation type, are the predominant upland vegetation over extensive areas 

of the Atlantic-Gulf-Caribbean coastal plain from the southeastern United States 

to Nicaragua and in parts of the West Indies (Robertson 1955, Luckhoff 1964). 

Within this geographical array, the "pineyard" of the northern Bahamas is by far 

the closest ecological counterpart of southern Florida pinelands. In both areas, 

pine forest occurs on a substrate of eroded limestone and has a species-rich 

understory of shrubby hardwoods and palms. The Bahaman pine forests - principally on the islands of Grand Bahama, Great Abaco, New Providence, and Andros -

are roughly twice as extensive as the original pine areas of southern Florida 

considered here and the woody flora of the Bahamas is considerably richer in 

species than that of southern Florida. To judge from brief habitat notes in the 

species accounts of floras of the Bahamas (Britton and Millspaugh 1920, Correll and 

Correll 1982), overall diversity of the shrub stratum in Bahaman pine forests 

appears to be somewhat greater than that recorded for all southern Florida pine 

areas combined (Table 6). N. L. Britton and C. F. Millspaugh noted 47 hardwoods 

as occurring in "pine-lands" or "pine-barrens" and Correll and Correll made similar 

mention of approximately 90 taxa in introductory comments 0982:23) or in text. 

The combined lists include about 95 taxa of hardwoods, palms, and agaves, and 
34 

presumably the plants listed were considered to be those more or less characteristic of the pineland understory. A list compiled according to the criteria used 

here for southern Florida, that is including species of infrequent or local occurrence in pineland, probably would show that at least 110 to 130 species enter the 

shrub stratum of Bahaman pine forests. Of the species listed, less than one-third 

(28 of c. 95) also occur in the shrub understory of southern Florida pinelands. As in 

southern Florida, most of the hardwood shrubs in Bahaman pine forests are also 

found (often as trees) in the "coppices", hardwood forests ecologically equivalent to 

southern Florida hammocks. The aspect of the understory in Bahaman pinelands 

differs principally in that palms tend to be much less prominent than in southern 

Florida. Serenoa is not known from the Bahamas. Thrinax, Coccothrinax, and. 

Sabal occur at least locally in the pineland shrub stratum, but palms do not 

commonly form either a distinct small-tree understory as in the Lower Florida 

Keys, or relatively continuous stands of palmetto-sized plants. 

Finally, one may note that the occurrence of pine forest on limestone in southern 

Florida and the northern Bahamas is unusual and apparently unexplained. Over its 

wide range, the tropical-subtropical, long-needle pine forest reportedly is found on 

sand, clay, gravel, shale, sandstone, granite, and other substrates, but not (or 

seldom) elsewhere on limestone. Almost throughout, local hardwood forest types 

occupy the limestone areas and pine forests commonly have abrupt interfaces with 

hardwood forest where limestone occurs, as in western Cuba (Henderson 1916, 

Seifriz 1943, Luckhoff 1964) and Belize (Russell 1964 and references cited there). 

The anomalous predominance of pine on limestone in southern Florida and the 

northern Bahamas invites the attention of ecologists. 

ACKNOWLEDGEMENTS 

Dr. Lloyd Loope was instrumental in starting the work for chapter 1 and provided 

the section on "fire history of hammocks." George Avery made species lists 

available, helped in the field surveys, and identified plants. Nancy Urban, Bob 

Russell, and Ron Bruckbauer helped with the elevation survey and vegetative 

analysis. In assembling data for chapter 2, the assistance of George Avery was, 

again, invaluable and Alan Herndon, Lance Gunderson, and Regina Rochefort 

provided helpful comment. Dee Childs, Jessie Brundige, and Betty Curl typed 

several manuscripts. We are grateful to them all. 
35 

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37 

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38 

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Table 1. Ground water levels (cm)* in pineland wells. 

1979 

Date Osteen Hammock 

5/9 

5/29 

6/18 

6/26 

7/17 

8/29 

9/18 

9/26 

9/27 

10/18 

10/28 

10/30 

67 

121 

99 

127 

132 

118 

100 

115 

123 

2/18 183 

2/23 105 

2/27 117 

3/22 158 

3/24 158 

4/13 188 

August, during Tropical Storm 

Dennis, after 17 inches rain 60 

9/20 100 

1981 

Redd Hammock 

33 

59 

36 

60 

58 

56 

40 

48 

41 

30 

57 

60 

2 

48 

41 

* Water levels were measured from the top of the well (rock surface) to the water 

surface. 
42 

Table 2. Summary of Vegetation Analysis 

a. Number of woody, forb, and graminoid species per vegetation type 

Transect 

Hammock 

Pineland 

Glades 

Total 

208 

51 

167 

114 

Woody* 

30 

49 

20 

. . . . . . . . . . . . . . . . . . . . . 

Hammock-pineland 

Pineland-glade 

Hammock-glade 

29 common species 

93 common species 

9 common species 

Forbs 

20 

90 

68 

Graminoids 

1 

28 

26 

b. Cover percentages for woody, forb, and graminoid species per vegetation type 

Hammock 

Pineland 

Glades 

Woody % 

15** 

21 

2 

c. Characteristics of pine distribution 

Mean density 

250-660/ha 

mean dbh 

14-15 cm 

* Includes woody vines. 

Graminoids % 

5 

11 

. mean height 

12-15 m 

Forbs % 

1 

6 

2 

mean basal arZa 

44-116 m /ha 

Total % 

16 

32 

15 

** The cover was estimated for shrub vegetation only and does not include canopy 

cover. Explanation in text. 
43 

Table 3. Mean cover percentage (per m

2

) and frequency of woody, forb, and graminoid species 

in each vegetation type. 

Pineland Hammock Glade 

(21 plots) (5 plots) (6 plots) 

Species Cover Frequency Cover Frequency Cover Frequency 

Woody 

Ardisia escallonioides .7 27 1.6 60 

Bumelia reclinata .1 8.5 .6 11.7 

B. salicifolia .8 23.3 .1 18 

Bursera simaruba • 1 8 

Byrsonima lucida .8 9.5 .4 3.2 

Chr~sobalanus icaco .3 9.5 .3 3.2 

Coccoloba diversifolia 2.5 4.4 

Conocarpus erectus .3 7.6 

Dodonoea viscosa 2.0 38.1 

Eugenia axillaris .1 .4 1.1 56 

Guettarda elliptica 1.1 18.1 

G. scabra 4.8 61.9 • 1 4 

Hex cassine .2 5.2 T 3.3 

Lysiloma 1atisiliquum .1 1.4 T 2 

Metopium toxiferum .9 15.7 .4 32 

Myrica cerifera .1 8.6 T 

Myrsine floridana .7 27.6 .6 34 

Persea borbonia .2 5.2 • 1 1.6 

Pinus elliottii .6 26.2 T 10 

Psychotr ia nervosa .1 .4 6 92 

Quercus virginiana .1 1.9 T 6 

Randia acu1eata .8 43.3 • 1 4.8 

Rhus copallina .9 18.1 

Serenoa repens 2.2 21.4 

Tetrazygia biflora .5 8.6 • 1 6 

Toxicodendron radicans • 1 6 

Trema micranthum .1 .9 

Sabal palmetto 1.3 23.8 T 2 

Forbs 

Aca1~pha chamaedrifolia T 1.9 

Adiantum tenerum .7 12 

Agalinis purpurea .1 10.0 T 3.3 

Aletris bracteata .1 2.9 T 3.2 

Anemia adiantifolia .5 55.2 • 1 4 

Angadenia sagraei .11 27.6 T 4.8 

Aster adnatus .1 4.8 T 21.0 

A. dumosus .1 10.4 T 8.3 

A. tenuifolius T 1.6 
44 

Table 3 continued. 

Pineland Hammock Glade 

(21 plots) (5 plots) (6 plots) 

SEecies Cover Freguencl:: Cover Freguencl:: Cover Freguencl:: 

Borreria terminalis .1 28.1 

Cassl::tha filiformis .1 19.0 T 6.6 

CenteHa asiatica .2 3.0 • 1 57.0 

Chamaesl::ce Einetorum .1 17.1 T 11.6 

C. Eorteriana Var. Eorter . I 29 T 18.0 

Chiococca Earvifolia .6 59.5 

Cirsium horridulum T 5 T 3.0 

CoreoEsis leavenworthii T 3.8 T 1.7 

Cynanchum blodgettii T 1.0 T 1.5 

Cl::noctenum mitreola T 5.2 T 25.0 

Dyschoriste oblongifolia .1 38.1 T 53.3 

EUEator ium leEtoEhl:: Hum .1 7.6 .1 15.0 

E. mikanioides T .5 T 6.6 

E. villosum .2 4.8 

Evolvulus sireceus T 7.1 T 40.0 

Galactia spp. T 0.9 T 2.0 

HeliotroEium 

Eoll::Ehyllum .1 11.9 • 1 43.0 

Hl::Etis alata yare 

stenoEhl:: lla .1 20.5 • 1 35.0 

IEomoea sagittata T .9 • 1 25.0 

Linu m medium .1 65.0 

Lobelia glandulosa T 1.9 T 3.3 

Melanthera angustifolia .2 53.3 

Mikania scandens . I 18.6 .1 26.6 

Mor inda rOl::oc .7 41.4 . I 8 

Passiflora suberosa .1 24.3 .1 4 · I 

Phl:: la nodiflora .1 0.9 · I 20 

Phl::llanthus caroliniensis T 0.5 T 4.9 

f. EentaEhl::llus .1 47.6 T 16.6 

Physalis viscosa .1 10.5 • 1 5 

Pir igueta caroliniana · T 7.6 T 10 

Pluchea rosea .1 7.6 .1 28.3 

Poll::gala .b0l::kinii T .5 T 1.7 

P. grandiflora T 2.9 T 10 

Pter idium aguilinu m 

yare caudatum .6 18.1 

Pter is longifolia var. 

bahamensis .4 26.7 

Ruellia caroliniensis .1 17.6 • 1 21.6 

Samolus ebracteatus .1 13.3 T 15 

Sida elliotii .1 2.9 • 1 13.3 

Smilax auriculata .2 16.7 .1 14 

Solidago stricta T 13.3 T 18 

Stylanthes calcicola T 2.8 .1 26 
45 

Table 3 continued. 

Pineland Hammock Glade 

(21 plots) (5 plots) (6 plots) 

Species Cover Frequency Cover Frequency Cover Frequency 

Thelypter is kunthii T 1 • 1 2 

Vernonia blodgettii T 4.3 T 26.6 

Vitis rotundifolia • 1 15.2 T 6 T 1.7 

Zamia pumila .1 9.5 .1 4 

Graminoids 

Andropogon cabanisii 2.5 62 

Andropogon glomeratus .1 8 .2 23 

A. virginicus T .9 T 4.8 

Aristida puq:~urascens .1 12 .2 70 

Cladium jamaicense .3 20 3.0 95 

Dicanthelium 

dichotomum T 4.3 .3 61.6 

Dichromena flor idensis .2 20.5 .4 53 

Muhlenbergia filipes .4 40 10.0 95 

Rhxnchospora divergens T 1.9 .2 66.6 

R. microcarpa T 5 • 1 31.6 

Schizachxr ium 

rhizomatum .3 21.4 .2 48.3 

S. semiberbe .3 14.3 T 1.7 

Setaria geniculata • 1 28.3 

Sorghastrum secundum .2 2.9 

T = trace. 
Table 4. Frequency and percent cover of certain tree and grass species (10 m2 plots) indicating distribution in high and low elevation pineland sites (explanation in text). 

3* .5 1 7 69 6.5 61 63 67 19 23 27 2.5 29 37 39 41 4.5 49 47 .53 

(1)** (2) (3) (4) (20) (18) (16) (I 7) (19) (.5) (6) (8) (7) (9) (10) (II) (12) (13) (I.5) (14) 

Lysiloma latisiliguum 30 

.5 

Guettarda scabra 60 60 100 100 90 90 100 100 100 10 10 10 100 90 100 90 100 

l.5 32 69 119 69 43 &7 101 3.5 2 2 T 109 62 14.5 46 106 

Guettarda elliptica 20 40 70 60 90 20 60 20 20 

3 6 13 12 127 2 67 .5 2 

Dodonea viscosa 100 70 80 40 10 30 20 10 40 100 90 10 10 100 8 10 

34 43 32 22 3 6 .5 2 2.5 99 23 T 4 8.5 4.5 3 

Metopium to-,~Jierum 10 30 90 .50 20 10 10 40 20 20 30 40 60 

T 2 .53 .5 T 2 10 28 6 2 10 7 74 

Bumelia salicifolia .50 30 10 40 10 .50 20 .50 10 100 20 60 

21 .5 T 27 1 13 2 l.5 2 60 9 7 

Randia aculeata 20 20 10 20 100 .50 60 20 60 .50 90 60 .50 30 60 

T .5 T T 61 3 22 4 12 9 17 & 11 11 .5 

Ardisia escaUonioides 70 70 10 70 40 20 10 40 30 20 20 40 70 60 

19 16 3 17 10 .5 1 .5 22 8 2 " 37 6 

~cassine 40 10 30 

II 1 14 

Rhus copallina 20 .50 10 40 80 100 80 

2 18 T 9 36 94 3.5 

~ 

a-. 
Table 4 continued. 

3* , 1 7 69 6.5 61 63 67 19 23 27 2.5 29 37 39 41 4.5 49 47 .53 

(1)** (2) (3) (4) (20) (18) (16) (17) (19) (.5) (6) (8) (7) (9) (10) (lJ) (12) (13) (I') (14) ---------~------ ------

Persea borbonia 20 60 30 

1 34 10 

Myrica cerifera 10 20 20 10 10 10 10 20 

6 1 T T T T T T 

ChrY5lObaianus icaco 10 30 20 10 40 

T 6 9 1 4 

Conocarpus ~ 40 30 40 30 60 

32 9 20 12 1) 

BumeUa reclinata 10 40 20 10 10 20 40 

1 3 1 T T 3 2 

Byrsonima lucida 10 10 '0 10 40 .50 20 10 

10 3 87 1 7 .50 2 6 

Andro~ cabanisii .50 20 40 10 100 100 100 90 100 60 100 70 100 100 90 70 100 

13 4 27 T 39 37 37 8.5 86 10 2.5 21 79 14 29 1.5 1.5 

Muhlenbergia filipes 100 70 100 100 100 100 100 .50 10 70 30 10 

6.5 3 111 47 33 .5S 6 6 T 29 T T 

Schizachyrizum rhizomatum 20 .50 70 60 80 )0 100 30 10 

2 9 12 8 14 .5 14 1 T 

• ActUilI plot numbers of Figure 1 • 

•• Numbers in parenthesis are numbers on ordination Figure 3. 

Top number = frequencf. 

Bottom number = cover percent. 

+=-

'-I 
48 

Table 5. Fire impact status and inferred fire history of hammocks, by year, as 

determined from aerial photography and helicopter flights in 1981. 

II Name 1940 1952 1960 1964 1980/81 

1 Twin Glades B M D for agriculture 

2 Barnes B M D for agriculture 

3 Torre M M M M M 

4 Gould M M M M 

5 DaB B R 1M 1M M 

6 Emery B B PM 

7 Lermond B R R 

8 Bequaert B R R M 

9 Simmons 1M M M M 

10 Gifford M B* R M 

11 Henderson M B R M 

12 No name M M 

13 No name B M 

14 No name B B/R M 

15 No name B M 

16 No name B B/R 1M 

17 No name B 

~ M 18 No name M R 

19 Rafinesque R B M 

20 Baker B 1M M 

21 Mosier M 

~ M M M 22 Rattlesnake 

~R R M 23 Osteen BNE R R M 

24 Gun R B R M 

25 Bartsch M M 

26 Palma Vista 112 M M M M 

27 Palma Vista 111 M M M M 

28 Smith (Small) M M M M 

29 McGinty R M M M 

30 Walton R B M M 

31 Deckert R B R PM 

32 Tomlin R R M ~E M 

33 Chase B R 1M 1M 

34 Turkey B R R R/IM M 

34A Mystery (Turkey) B R M M 

35 Allen B R R PM 

36 Marshall B BNW M 

37 Clapp B B R M 

38 PHsbry B M 

38A Redd M M M M M 

40 Goodrich M B** R M 

41 Johnson R BSW R M 

42 Fairchild B B** R PM 

43 Archer R M M M 

\ 
Table 5 continued. 49 

II Name 1940 1952 1960 . 1964 1980/81 

44 Lea B R R 1M 

45 Conrad 1M 

46 Call B*** R R 1M M 

47 DeCamp B 

~ R R M 48 Currier B R R M 

49 Rehder B B BN R 

50 Bootlegger M M B 1M 

51 Dark M M M partial recent burn 

52 Simpson B B B R R 

53 Gill B B B R M 

54 Cadwa1eder B BE R M 

55 Barbour R B M 

56 Say B R M M M 

57 Wright 

~ M M M M 58 Winkley R M M M 

59 Clench M B R R M 

60 Frampton B R R R M 

61 No name R B M 

62 No name B R M 

63 Pfleuger R 1M M 

64 Warren B R 

65 Lime M M 

66 Squires M R 

67 West End R M 

68 ScheviU M B,R 

69 McDonald R M M M 

70 Poppenhager B B B BS M 

71 Lott B  R 1M B M 

72 Dr. deBoe B R R R 

73 Top B B R R BS , M 

76 Junk 1M M M M M 

77 Jackson B R R PM 

79 Pineridge B B B M 

80 Clavata B R,BW R 1M 

81 Rhodes D 

85 Fields R 1M M R 1M 

86 Maxwell B R R R 1M 

87 Mathew M 

88 No name R B R M 

89 Dead B R 1M M 

90 Barret Bw R R R M 

91 Deer B 

~ M M 1M 

92 Von Paulsen M R M M 

93 Grimshawe M M M M M 

94 Winkleman B B R R M 

95 Coe M B,R R M M 

96 Powell ~BSE R 

97 Remington B +E M 

98 Tryon B 

99 Humes B B**  R M 

n 
50 

Table 5 continued. 

II Name 1940 1952 1960 1964 1980/81 

100 Beard B B - R1969 M 

101 Pennecamp B R BNW , R M 

102 Jones M B R M M 

104 Craighead M BW R M M 

105 Wild lime M B R M M 

106 Ebbitts B R R R M 

107 Robertson (A - F) B R M 

108 ' White rock B B B B 1M 

109 Lysiloma B,R R M 1M 

110 F. N. Young B R R M 

111 Simpson B,R B R M 

112 Abbott M B R M 

113 Buttonwood M 

114 No name M 

115 No name M 

116 Avery M 

117 Douglas (Mar jor ie Stoneman) M 

118 Simmons (Glenn and Maxie) 

119 Young, Fran M 

120 Brookfield M 

Legend: 

Fire Impact Status 

M = Hammock has mature appearance on aerial photograph 

PM = Hammock has mature appearance on aerial photograph, except for small shrubby 

portions. 

1M = Hammock appears to be in condition approaching maturity on aerial photograph. 

R = Hammock exhibits signs of progressive recovery from burn in previous interval(s) 

D = Destroyed 

B = Evidence of burn during interval (N, S, W = north side, south side, etc.) (Mar = margins) 

* scattered tall trees 

** center 

*** part 

**** understory 
51 

Table 6. Distribution of 95 taxa of shrub-layer plants in pine forest areas of southern 

Florida. 

So. No. 

Shrub Species LFK LPK Biscyane Biscayne BICY 

Acacia farnesiana x 

x (4)2 

x 

Acacia pinetorum x x 

Alvaradoa amorphoides x x 

Amorpha crenulata x 

Annona glabra x x (7) x x 

Ardisia escallonioides x x (93) x x 

Asimina reticulata x 

Bacchar is angustifolia x 

Baccharis glomeruliflora x (14) x x 

Bacchar is halimifolia x x (14) x x x 

Befar ia racemosa x 

Bourrer ia cassinifolia x x (1) x x 

Bumelia celastrina x x 

Bumelia reclinata x (64) x x x 

Bumelia salicifolia x x (96) x x x 

Bursera simarOba x x (8) x 

Byrsonima lucida x x (86) x x 

Caesalpinia pauciflora x 

Callicarpa americana x (4) x x x 

Cassia chapmanii x x (34) x x 

Catesbaea parviflora x 

Cephalanthus occidentalis x (1) x 

Chrysobalanus icaco x x (71) x x x 

Chrysophyllum oliviforme x x (44) 

Citharexylum fruticosum x (10) x 

Coccoloba diversifolia x x (10) 

Coccoloba uvifera x 

Coccothrinax argentata x x,(18) x x 

Colubrina arborescens x 

Colubrina cubensis x 

Conocarpus erectus x x (22) x 

Crossopetalum rhacoma x x x 

Croton linear is x x (42) x x 

Diospyros virginiana x x 

Dodonaea viscosa x (64) x x x 

Drypetes diversifolia x 

Erithalis fruticosa x 

Eugenia axillaris x x (53) x x 

Eugenia myrtoides x 

Eupatorium villosum x (26) x 

Exothea paniculata x (10) x 

Ficus aurea x x (5) x x x 

Ficus citr ifolia x x (62) x x x 

= 
52 

Table 6 continued. 

So. No. 

Shrub Species LFK LPK Biscyane Biscayne BICY 

Forestiera segregata x (16) x 

Guapira discolor x x (71) x 

Guettarda elliptica x x (77) x 

Guettarda scabra x x (52) x x 

1 Hippomane mancinella x 

Hypelate tr ifoliata x x 

llex cassine x (82) x x x 

Hex glabra x 

Hex krugiana x (27) x 

Jacguinia keyensis x x 

Laguncular ia racemosa x 

Lantana involucra ta x x (21) x x 

Lyonia fruticosa x x 

L ysiloma latisiliguum x (8) x 

Magnolia virginiana x 

Manilkara bahamensis x 

Mastichodendron foetidissimum x (3) 

Metopium toxiferum x x (97) x x 

Myrcianthes fragrans x (4) x 

Myrica cerifera x x (93) x x x 

Myrsine floridana x x (99) x x x 

Per sea borbonia x x (74) x x x 

Piscidia piscipula x 

Pisonia rotundata x 

Pithecellobium guadalupense x 

Psidium longipes x x (67) x x 

Psychotr ia nervosa x x (4) x 

Quercus minima x 

Quercus pumila x 

Quercus virginiana x (23) x x x 

Quercus va. var. geminata x x 

Randia aculeata x x (96) x x x 

Reynosia septentr ionalis x 

Rhizophora mangle x 

Rhus copallina x x (44) x x x 

Sabal palm etto x x (100) x x x 

Salix caroliniana x x x x 

Sambucus simpsonii x 

Serenoa repens x (92) 

Solanum donianum x x 

Solanum erianthum x 

Sophora tomentosa x 

Strumpfia maritima x 

Sur iana mar itima x 

Taxodium ascendens x 

Tetrazygia bicolor x (90) x x 

Thr ina x morr issii x 
• 

Table 6 continued. 

Shrub Species 

Trema lamarckianum 

Trema micranthum 

Vaccinium myrsinites 

Ximenia americana 

LFK = Lower Flor ida Keys. 

LPK = Long Pine Key. 

LFK LPK 

x x (5) 

53 

So. No. 

Biscyane Biscayne BICY 

x 

x x 

x x 

x 

I Neither Small (1933:829) nor Long and Lakela (1976:574) mentions occurrence of this 

species on the Florida mainland, but it is frequent in a relatively small area of 

sou thwestern Long Pine Key (Robertson 1953, Figure 23; South Flor ida Research Center 

Herbarium Nos. 49 and 2199). Our listing for the Lower Florida Keys is based on Small 

(loc. cit.). We have not seen the species there in pineland situations. 

2 () percentage of constancy of woody species encountered in 73 plots (see pineland shrub 

methods, Chapter 1) • 
54 

Table 7. Summary: shrub stratum diversity and similarity in southern Florida pine forests. 

No. Unique No. of Taxa in Common 

Area No. Taxa Taxa 1 2 3 4 5 

1. Lower Florida Keys 59 20 38 32 24 14 

2. Long Pine Key 61 5 38 49 31 21 

3. Southern Biscayne 49 0 32 49 31 20 

4. Northern Biscayne 41 6 24 31 31 21 

5. Big Cypress 28 4 14 21 20 21 

Table 8. Density of shrub-stratum plants in southern Florida pine forests. 

Area No. Plants/100 m 

2 Percent Hardwoods Percent Palms 

Big Pine Key 1 400 85 15 

Long Pine Key I 919 96 4 

1 (360-1470) (89-100) (0-11) 

Southern Biscayne 1290 97 3 

(Navy Wells) 1 Southern Biscayne 615 83 17 

(Camp Owaissa qauer) 

Northern Biscayne 695 85 15 

(Thompson Park)1 

Northern Biscayne 605 70 30 

(USDA) 1 Northern Biscayne 265 81 19 

(Tamiami Pi~s) 

Long Pine Key 149 

(57-313) 

1 Calculated from Loope et al. (1979, Table 1). Numbers represent number of ideptifiable 

individuals of all sizes. Extrapolated from counts on a series of 20 plots each/m in each 

area. Long Pine Key data are the mean and range for seven such series of plots. 

2 From Taylor and Herndon (I981, Tables 1 and 2). Numbers represent hardwood stems 

3 feet(0.9 m) or more high. Reduced from data given as no. stems/hectare. 
, 

:J 400 

(J) 

:E 

-

~ 300 

(.) 

Z 

-

z 

Q 200 

~ > LLJ 

c ~ C 

Z (.) Z 

<t 0 <t LLJ -.J ~ -.J C LLJ ~ LLJ <t z <t Z .-.J a: - :z: 0- (!) 

C C 0 ~ Z Z Z (.) 

<t LLJ <t LLJ <t 0 --l C -.J C -.J :E LLJ <t LLJ <t LLJ :E Z -.J Z -.J Z <t a: - a: (!) 0- (!) :z: 

..J I 1/ / / / / / / / / / / / 4 / / / / / / / / / / / / I) / / / / / / / / / / / / /1/ / / / / / / / /. 

IJ.J 100 0 600 1200 1800 

E 

F7l 8iJ 

~ 

ORGANIC SOl L 

MARL 

LIMESTONE 

DISTANCE IN METERS w 

Figure I Elevation transect from Osteen to Redd Hammock {Number 1-71 

are plot numbers}. 

VI 

VI 
x 

w 

c 

z 

z 

o 50 

~ Z 

-

c 

a:: 

o 

*3 

2*1 

*4 

114*15 20 

6**:11 17 19 

8* 5 18* *12 

*13 

10/

16 

*9 

*11 

22* *21 

*26 

24* .-125 

23 o~ ~ 

o 

ORDINATION INDEX 

Figure 2 Ordination of 3 vegetation types: 

1, 2, 3, 4 - Hammock plots. 

5-20 - Pineland plots .• 

21-26 - Prairie plots. 

Equivalence of Figure 2 numbers to 

actual plot numbers on elevation 

transect (Fig. 1), first number is from 

ordination figure: 

1 - 9 

2 - 13 

3 - 15 

4 - 71 

5-3 

6-5 

7-7 

8 - 25 

9 - 27 

10 - 29 

11 - 37 

12 - 39 

13 - 41 

14 - 45 

15 - 47 

16 - 49 

17 - 61 

18 - 65 

19 - 67 

20 - 69 

21 - 21 

22 - 33 

23 - 55 

24 - 57 

25 - 59 

26 - 51 

\II 

0\ 
--

x 

l1J 

C 

z 

~ 50l 

I j 

-i 

.:1 2 *1 

*3 

o 1. 

0 

* 10 

9 

**8 

II 

* * *5 

16 *7 12 

***17*18 

6 *14 

* * *19 13 

20 

,4 I 

50 

OR DINATION INDEX 

* 15 

Figure 3 Ordination of pineland stands 

Equivalence of Figure 3 numbers 

to actual plot numbers of 

Figure 1: 

1 - 3 

2-5 

3 - 1 

4-7 

5 - 19 

6 - 23 

7 - 25 

8 - 27 

9 - 29 

10 - 37 

11 - 39 

12 - 41 

13 - 45 

14 - 47 

15 - 49 

16 - 61 

17 - 63 

18 - 65 

19 - 67 

20 - 69 

VI 

"-.I 
58 

Appendix I. Numbers and names of Long Pine Key hammocks. Asterisk indicates named 

and numbered hammocks not found on vegetation map. Names and/or 

numbers in parentheses indicate synonymous hammocks. 

1 Twin Glade 

2 Barnes 

3 Torre 

4 Gould 

5 Dall 

6 Emery 

7 Lermond 

8 Bequaert 

9 Simmons 

10 Gifford 

11 Henderson 

12 No name 

13 No name 

14 No name 

15 No name 

16 No name 

17 No name 

18 No name* 

19 Rafinesque 

20 Baker 

21 Mosier 

22 Rattlesnake 

23 Osteen 

24 Gun 

25 Bartsch 

26 Palma Vista 112 

27 Palma Vista III 

28 Smith (Small) 

29 McGinty 

30 Walton 

31 Deckert 

32 Tomlin 

33 Chace 

34 Turkey (72-deBoe, 

74-Gold, 75-Lowe) 

34A Mystery (34-Turkey) 

35 Allen 

36 Marshall 

37 Clapp 

38 Pilsbry 

38A Redd 

39 Winslow* 

40 Goodrich 

41 Johnson 

42 Fairchild 

43 Archer 

44 Lea 

45 Conrad 

46 Call 

47 DeCamp 

48 Currier 

49 Rehder 

50 Bootlegger 

51 Dark 

52 Simpson 

53 Gill 

54 Cadwalader 

,o5 Barbour 

56 Say 

57 Wright 

58 Winkley 

59 Clench 

60 Frampton 

61 No name 

62 No name 

63 Pfleuger 

64 Warren 

65 Lime 

66 Squires 

67 West End* 

68 Schev ill * 

69 McDonald 

70 Poppenhager 

71 Lott 

72 Dr. deBoe (34-Turkey) 

73 Top 

74 Gold (34-Turkey) 

75 Lowe (34-Turkey) 

76 Junk 

77 Jackson 

78 Sisal* 

79 Pineridge 

80 Clavata 

81 Rhodes 

82 No name* 

83 No name* 

84 Weber 

85 Fields 

86 Maxwell 

87 Matthew 

88 No name 

89 Dead 

90 Barret 

91 Deer 

92 Von Paulsen 

93 Grimshawe 
59 

Appendix I continued. Numbers and names of Long Pine Key hammocks. 

94 Winkleman 

95 Coe 

96 Powell* 

97 Remington* 

98 Tryon* 

99 Humes 

100 Beard 

101 Pennecamp 

102 Jones 

103 Winte* 

104 Craighead 

105 Wild lime 

106 Ebbitts 

107 Robertson 

108 White rock 

109 Lysiloma 

110 Young 

111 Simpson 

112 Abbott 

113 Buttonwood 

114 No name 

115 No name 

116 Avery 

117 Douglas (Marjorie Stoneman) 

118 Simmons(Glen and Maxie) 

119 Young (Fran) 

120 Brookfield 
60 

Appendix II. Species List 

Hammock Pineland Glade 

Acacia pinetorum x x 

Acalypha chamaedrifolia x 

Adiantum tenerum x 

Agalinis purpurea x x 

Aletris bracteata x x 

Andropogon cabanisii x 

Andropogon glomeratus x x 

Andropogon virginicus x x 

Anemia adiantifolia x x 

Anemia wrightii x 

Angadenia sagraei x x 

Annona glabra x 

Ardisia escallonioides x x x 

Aristida purpurascens x x 

Asclepias lanceolata x 

Asclepias longifolia x 

Aster adna tus x x 

Aster dumosus x x 

Aster tenuifolius x 

Ateramnus lucida x 

Baccharis glomeruliflora x x 

Baccharis halimifolia x x 

Borreria terminalis x 

Bourreria cassinifolia x 

Buchnera flor idana x x 

Bumelia reclinata var. reclinata x x 

Bumelia salicifolia x x 

Bursera simaruba x 

Byrsonima lucida x x 

Calyptranthes pallens x 

Callicarpa americana x 

Cassia chapmanii x 

Cassia deeringiana x 

Cassytha filiformis x x 

Catopsis berteroniana x 

Centella asiatica x x 

Centrosema virginianum x 

Cephalanthus occidentalis x 

Chamaesyce adenoptera x 

Chamaesyce pinetorum x x 

Chamaesyce porteriana x x 

Chiococca alba x 

Chiococca parvifolia x x 

Chrysobalanus icaco x x 

Chrysophyllum oliviforme x x 

Cirsium horridulum x x 
61 

Appendix II. Species List continued. 

Hammock Pineland Glade 

Citharexylum fruticosum x x 

Cladium jamaicense x x 

Coccoloba diversifolia x x 

Coccothrinax argentata x 

Coelorachis rugosa x 

Conocarpus erecta x x 

Coreopsis leavenworthii x x 

Crossopetalum ilicifolium x 

Crotalaria pumila x 

Croton linearis x 

Cynanchum blodgettii x x 

Cynoctenum mitreola x x 

Desmodium lineare x 

Dichanthelium dichotomum x x 

Dichanthelium spp. x x 

Dichromena colorata x 

Dichromena flor idensis x x 

Digitaria pauciflora x x 

DiocHa virginiana x 

Dodonaea viscosa x 

Dyschoriste oblongifolia x x 

Echites umbellata x 

Elytraria caroliniensis x 

Encyclia tampensis x x 

Eragrostis elliottii x x 

Eriochloa michauxii x 

Erythrina herbacea x 

Eugenia axillar is x x 

Eupatorium coelestinum x x 

Eupator ium leptophy Hum x x 

Eupator ium mikanioides x x 

Eupatorium villosum x 

Eustachys glauca x x 

Eustachys petraea x x 

Evolvulus sericeus x x 

Exothea paniculata x 

Ficus aurea x 

Ficus citrifolia x x 

Forestiera segregata var. pinetorum x 

Fuirena brev iseta x 

Galactia spiciformis x 

Galactia spp. x x 

Galium hispidulum x 

Guapira discolor var. longifolia x x 

Guettarda elliptica x 

Guettarada scabra x x 

Hedyotis nigricans var. filifolia x 
62 

Appendix II. Species List continued. 

Hammock Pineland Glade 

Heliotropium polyphyllum x x 

Heterotheca graminifolia var. tracyi x 

Hymenocallis palmeri x 

Hypericum brachyphyllum x 

Hypericum hypericoides x x 

Hyptis alata var. stenophylla x x 

Hex cassine x x 

Hex krugiana x 

Ipomoea microdactyla x 

Ipomoea sagittata x x 

Ipomoea tenuissima x 

Iva microcephala x x 

Jacguernontia curtissii x 

Justicia ovata x 

Lantana depressa x 

Lantana involucrata x 

Lasiascis divaricata x 

Leipharmos parasitica x 

Linum medium x 

Lobelia glandulosa x x 

Ludwigia microcarpa x 

Ludwigia repens x x 

Lysiloma latisiliguum x x 

Mastichodendron foetidissimum x 

Mecardonia acuminata x 

Melanthera angustifolia x 

Melanthera parvifolia x 

Metopium toxiferum x x x 

Mikania scandens x x 

Mor inda royoc x x x 

Muhlenbergia filipes x x 

Myrcianthes fragrans x 

Myrica cerifera x x 

Myrsine floridana x x x 

Nectandra coriacea x 

Oxypolis filiformis x x 

Panicum hemitomon x 

Panicum rigidulum x x 

Panicum tenerum x x 

Panicum virgatum yare cubensis x x 

Parthenocissus guinguefolia x 

Paspalum caespitosum x 

Paspalum monostachyum x 

Paspalum setaceum x 

Passiflora suberosa x x x 

Persea borbonia x x 
63 

Appendix II. Species List continued. 

Hammock Pineland Glade 

Phyla nodiflora x x 

P. stoechadifolia x x 

Phy llanthus caroliniensis x x 

Phyllanthus pentaphyllus var. floridanus x x 

Physalis viscosa x x 

Pinus elliottii var. densa x x 

Pisonia aculeata -- x 

Pirigueta caroliniana var. glabra x x 

Pirigueta caroliniana var. tomentosa x x 

Pluchea rosea x x 

Poinsettia pinetorum x 

Polygala balduinii x 

Polygala boykinii var. sparsifolia x x 

Polygala grandiflora x x 

Polypodium phyllitidis x 

Polypodium polypodioides x 

Proser pinaca palustr is x 

Prunus myrtifolia x x 

Psidium longipes x 

Psychotria nervosa x x 

P. sulzneri x 

Pteridium aguilinum var. caudatum x 

Pter is longifolia var. bahamensis x x 

Quercus virginiana x x 

Randia aculeata x x 

Phus copallina x 

Rhynchosia minima x 

Rhynchospora divergens x x 

R. microcarpa x x 

Ruellia caroliniensis spp. ciliosa x x 

Sabal palmetto x x 

Sabatia grandiflora x 

Sachsia polycephala x 

Samolus ebracteatus x x 

Schinus terebinthifolius x 

Schizachyrium gracile x 

Schizachyrizum rhizomatum x x 

Schizachyrium sanguineum x x 

Schoepfia chrysophy lloides x 

Scler ia ciliata x 

Serenoa repens x x x 

Setaria geniculata x x 

Setaria spp. x 

Sida elliottii x x 

Simarouba glauca x x 

Smilax auriculata x x ---

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64 

Appendix II. Species List continued. 

Hammock Pineland Glade 

Solanum donianum x x 

Solidago chapmanii x x 

Solidago str icta x x 

Sorghastrum secundum x 

Stillingia sylvatica ssp. tenuis x x 

Stenandrium dulce x 

Stylosanthes calcicola x x 

T ephrosia flor ida x 

Tetrazygia bicolor x x x 

Teucr ium canadense x 

Thelypteris kunthii x x x 

Tillandsia balbisiana x 

Tillandsia circinnata x 

Tillandsia fasciculata x 

Tillandsia valenzuelana x x 

Toxicodendron radicans x x 

Tragia saxicola x 

Trema micranthum x 

Trichomanes kraussi x 

Tripsacum floridanum x 

Vernonia blodgettii x x 

Vitis rotundifolia x x x 

Waltheria indica x x 

Zamia pumila x x