Distribution and character of natural habitats in pre-settlement northern Florida, as recorded by public land survey records

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Title:
Distribution and character of natural habitats in pre-settlement northern Florida, as recorded by public land survey records
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Book
Creator:
Schwartz, Mark W.
Travis, Joseph
Publisher:
Florida Game and Fresh Water Fish Commission
Place of Publication:
Tallahassee, Fla.
Publication Date:

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University of Florida
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University of Florida
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ltqf - AAA1620
ltuf - AME1729
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UF00000501:00001

Table of Contents
    Front Cover
        Page i
        Page ii
    Figures
        Page iii
    Main
        Figure 1
        Figure 2
        Figure 3
        Figure 4
        Figure 5
        Figure 6
        Figure 7
        Figure 8
        Figure 9
        Figure 10
        Figure 11
        Figure 12
        Figure 13
    List of Figures
        Figure 14
    Tables
        Table 1
        Table 1a
        Table 2
        Table 2a
        Table 3
        Table 4
        Table 5
        Table 6
        Table 6a
        Table 7
        Table 8
        Table 8b
        Table 9
        Table 9b
    Two
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
    Back Cover
        Page 23
        Page 24
Full Text














The .Distribution and Character-of Natural Habitats in

: sefttlemnt Northern Florida,
-as.Recordeid i Public Land Survey Records


PROJECT REPORT




Mark W. Schwartz
Joseph Travis.


December 1995


Nongame Wildlife Programi
Florida GC me and Fresh Water Fish Commission
620 South Meridian Street '
S-Tallahassee. FL32399-1600
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The Distribution and Character of Natural Habitats in
Pre-settlement Northern Florida,
as Recorded by Public Land Survey Records






Mark W. Schwartz1
Joseph, Travis2



'Center for Biodiversity
Illinois Natural History Survey
607 East Peabody Drive
Champaign, IL 61820
2Department of Biological Sciences
Florida State University
Tallahassee, FL 32306-2043










Submitted as project report for
Nongame Wildlife Program project GFC-86-020


December 1995


;IES




















-I-.


hence t these ere a0 e i in(
township (a) was surveydein 18 while (b) was survey n


~I:jii~


~~ ~t~~ylCch




Township


Samples


1 Mile





















































Figure 3. The distribution of pineland communities of northern
Florida as defined by Public Land Survey Records.



























































Figure 4. The distribution of mixed wetland communities of
northern Florida as defined by Public Land Survey Records.









Marv;























































Figure 5. The distribution of beech (Facus qrandifolia) of
northern Florida as defined by Public Land Survey Records.





































Figure 6. The distribution of cypress (Taxodium ascendens and
Taxodium distichum) of northern Florida as defined by Public Land
Survey Records.






- -- -

v~X I


















Figure 7. The distribution of cypress-dominated communities of
northern Florida as defined by Public Land Survey Records.


''.te &^"^^^L '^BB~t



































































Figure 8. The distribution of titi (Cvrilla racemiflora or
Cliftonia monophvlla) of northern Florida as defined by Public
Land Survey Records.










-- --- -- -






























Figure 9. The distribution of titi-dominated communities of
northern Florida as defined by Public Land Survey Records.





.... ... ...














































Figure 10. The distribution of gum-dominated (Nvssa sp.)
communities of northern Florida as defined by Public Land Survey
Records.







I ''



k .' 4T- "'






















































Figure 11. The distribution of bay-dominated (Magnolia
virginiana, Gordonia lasianthus, or Persea sp.) communities of
northern Florida as defined by Public Land Survey Records.


































Figure 12. The distribution of mixed hardwood wetland communities
of northern Florida as defined by Public Land Survey Records.







I!


14 h~~~Li
























































Figure 13. The distribution of cabbage pala (Sabal palmetto) of
northern Florida as defined by Public Land Survey Records.




FIGURE LEGENDS

Figure 1. A map of northern Floridaooutlining Public Land Survey
section marker vegetation records tabulated in this study. AV0k4,,a.

Figure 2. Two man of ownships ith maj l rladdmm s. These
townships are adj ce t on/ano e howe er, the 4 eys, and
hence these maps wer s etch at 30 ear /inter alsl.) Te first
township (a) was u d in 18 while ( wass" eyed in 18

Figure 3. The distribution of pineland communities of northern
Florida as defined by Public Land Survey Records.

Figure 4. The distribution of mixed a communities of
northern Florida as defined by Public Land Survey Records.

Figure 5. The distribution of beech (Faaus arandifolia) of
northern Florida as defined by Public Land Survey Records.

Figure 6. The distribution of cypress (Taxodium ascendens and
Taxodium distichum) of northern Florida as defined by Public Land
Survey Records.

Figure 7. The distribution of cypress-dominated communities of
northern Florida as defined by Public Land Survey Records.

Figure 8. The distribution of titi (Cyrilla racemiflora or
Cliftonia monophvlla) of northern Florida as defined by Public
Land Survey Records.

Figure 9. The distribution of titi-dominated communities of
northern Florida as defined by Public Land Survey Records.

Figure 10. The distribution of gum-dominated (Nyssa sp.)
communities of northern Florida as defined by Public Land Survey
Records.

Figure 11. The distribution of bay-dominated (Magnolia
virriniana, Gordonia lasianthus, or Persea sp.) communities of
northern Florida as defined by Public Land Survey Records.

Figure 12. The distribution of mixed hardwood wetland communities
of northern Florida as defined by Public Land Survey Records.

Figure 13. The distribution of cabbage palm (Sabal palmetto) of
northern Florida as defined by Public Land Survey Records.




Table 1. A list of 71 species sampled in the Plublic Land Survey
data for northern Florida. Names encountered in the survey
records fewer than 10 times were omitted from this table.


Common Name


Number of
occurences


percent of
sample


Pine
Oak
Cypress
Gum
Bay
Redbay
Sweetbay
Water Oak
White Oak
Magnolia
Hickory
Ash
White Bay
Beech
Cedar
Cherry
Chinquapin
Dogwood
Elm
Sweetgum
Black Gum
Tupelo Gum
Hackberry
Haw
Red Haw
Holly
Hornbeam
Ironwood
Lightwood
Linden, Linn
Locust
Maple
Mulberry
Myrtle
Black Oak
Post Oak
Spanish Oak
Swamp Oak
Blackjack Oak
Live Oak
Willow Oak
Red Oak
Cabbage Palm
Palmetto
Saw Palmetto
Persimmon
Shortleaf Pine
Longleaf Pine


101,065
964
3801
1535
6134
121
11
886
391
441
860
570
76
332
40
21
152
552
140
281
349
8190
24
42
17
316
110
152
187
44
10
354
10
147
472
582
16
58
3360
432
48
1511
400
72
36
70
19
137


77.015
0.735
2.897
1.170
4.674
0.092
0.008
0.675
0.298
0.336
0.655
0.434
0.058
0.253
0.030
0.016
0.116
0.421
0.107
0.214
0.266
0.678
0.018
0.032
0.013
0.241
0.084
0.116
0.143
0.034
0.008
0.270
0.008
0.112
0.360
0.444
0.012
0.044
2.560
0.329
0.037
1.151
0.305
0.054
0.027
0.053
0.014
0.104







Table 1. A list of 71 species sampled in the Plublic Land Survey
data for northern Florida. Names encountered in the survey
records fewer than 10 times were omitted from this table.

Common Name Number of percent of
occurences sample
------------------------------------------------------------------
56 Swamp Pine 57 0.043
57 Poplar 69 0.053
58 Privet 67 0.051
60 Sassafras 49 0.037
61 Sourwood 230 0.175
62 Whortleberry 34 0.026
64 Willow 75 0.057
65 Yaupon 31 0.024
67 Birch 12 0.009
71 Spruce (pine) 30 0.023
72 Titi 1784 1.359
76 Sumac 10 0.008
77 Spruce Pine 52 0.040
84 Can't Read* 51 0.039
85 Unusual Tree* 86 0.066
86 No Tree* 761 0.580
88 Pit* 29 0.022
89 Mound* 132 0.101
91 Juniper 17 0.013
92 Laurel 40 0.030
99 Cucumber Tree 16 0.012
107 Baygall 20 0.015
112 Dead* 15 0.011
Other trees 113 0.086

* Codes that indicate where corner marker was not an
identifiable tree




Table 2. A list of woody species found in north Florida and their
associated common names, as found in the Public Land Survey
records.


Common Name

GYMNOSPERMS


Cedar
Cypress

Pine


Latin Name


Chamaecyparis thvoides. Juniperus silicola
Taxodium ascendens, T. distichum

Pinus clausa. P. echinata, E. elliottii,
P. alabra. P. palustris. P. serotina. P. taeda


Longleaf Pine
White Pine
Shortleaf Pine
Swamp Pine
Spruce Pine
Spruce


P. palustris
P. palustris
P. echinata
P. elliottii. P. serotina
P. qlabra
P. glabra


ANGIOSPERMS


Ash Fraxinus americana. F. caroliniana.
F. pauciflora. F. pennsvlvanica. F. profunda
Bay Persea borbonia. P. palustris.
Magnolia virginiana. Gordonia lasianthus
Redbay Persea borbonia. P. palustris
Whitebay Magnolia virginiana
Bullbay Magnolia grandiflora
Beech Faaus grandifolia
Birch,Burch Betula nigra
Cherry Prunus americana. P. anqustifolia
P. caroliniana. P. serotina. P. umbellata
Chinquapin Castanea shei C. floridana
Cucumber Tree Magnolia oat
Dogwood Cornus alternifolia. C. florida. C. foemina,
Swamp Dogwood Cornus foemina
Elm Ulmus alata. U. americana, U. rubra
Red Elm Ulmus SD.
Grape Vine Vitis sp.
Gum Nvssa aquatica. N. biflora. N. oqeche,
N. sylvatica
Black Gum Nyssa biflora. N. sylvatica
Tupelo Gum Nvssa acuatica. N. biflora. N. oaeche,
N. sylvatica
Hackberry Celtis laevigata
Haw Cretaequs sPp.
Red Haw Cretaequs spathulata
May Haw Cretaecus aestivalis
Hickory (Hickrey Carva aguatica. C. cordiformis, C. floridana
C. glabra. C. ovalis, C. pallida, C. tomentosa
Holly Ilex opaca





Table 2. A list of woody species found in north Florida and their
associated common names, as found in the Public Land Survey
records.


Common name
----------------
Hornbeam
Ironwood
Juniper
Laurel
Linden, Linn
Locust
Magnolia
Maple

Mulberry
Myrtle

Oak
Black Oak
Blackjack Oak
Live Oak
Post Oak
Spanish Oak
Swamp Oak
Water Oak
White Oak
Willow Oak
Palm
Palmetto
Cabbage Palm
Saw Palmetto
Persimmon
Poplar

Privet
Redbud
Sassafras
Sourwood
Sumac (Shoemake)
Sweetbay
Sweetgum
Titi

Whortleberry
Willow


Latin Name


Carpinus carolinianum
Ostrva virciniana
Juniperus silicola
Ouercus hemisphaerica
Tilia americana
Gleditsia triacanthos
Magnolia grandiflora
Acer barbatum, A. leucoderme. A. rubrum
A. saccharinum
Morus alba, M. rubra
Myrica cerifera. M. inodora
Ilex myrtifolia. Ouercus myrtifolia
Quercus spp (25 species)
Q. velutina
Q_ marilandica
Q. virginiana (Q. hemisphaerica. Q. qeminata)
OQ stellata
Q. falcata
Q. prinus (Q. michauxii. Q. falcata)
Q. nigra
Q. alba
0. phellos
Sabal palmetto
S. palmetto (??)
S. palmetto
S. palmetto (??)
Diospyros virginiana
Populus deltoides. P. heterophylla
Liriodendron tulipifera
Foresteria acuminata
Cercis canadensis
Sassafras albidum
Oxydendrum arboreum
Rhus copallina
Magnolia virginiana
Liquidambar stvraciflua
Cyrilla parviflora. C_ racemiflora
Cliftonia monophylla
Ilex sap (??)
Salix caroliniana, S. floridana, S. niqra
















Table 3. A list of pine species in north Florida, and a brief
description of the habitats in which they are found (from Kurz
and Godfrey, 1962).


common name

(sand)


Pinus echinata (shortleaf)


Pinus elliottii (slash)



Pinus glabra (spruce)


Pinus palustris (longleaf)



Pinus serotina (pond)


Pinus taeda


(loblolly)


habitat


common near the Atlantic and Gulf
coasts on sand ridges, associated
with evergreen oaks

invades fertile upland old fields
and found in mixed hardwood stands

wet flatwoods, branch swamps,
along shallow ponds and bays near
coast

mixed hardwoods and hammocks,
and bottomland forests

uplands, flatwoods, and sand
ridges, occurs as dominant in
stands

poorly drained sites with marked
fluctuations in water table

invades upland old fields, and
mixed in lowland hardwoods


Species

Pinus clausa





Table. A list of community types identified in the PLS data for north
Florida, along with their abundances.


Community
Pine


Numb4


1.)
2.)
3.)
4.)
5.)
6.)
7.)
8.)
9.)
10.)
11.)
12.)
13.)
14.)
15.)
16.)
17.)
18.)
19.)
20.)
21.)
22.)
23.)


Total


Pine/oak
Pine/unresolved
Pine/oak/hickory
Pine/mixed hardwood
Pine/titi
Pine/Cypress
Pine/gum
Pine/bay
Pine/mixed wetland
Pine/other species
Pine/palm
Pineland associates
Oak/Hickory
Mesic Hardwoods
Titi swamp
Cypress mix
Gum swamp
Bay swamp
Mixed wetland
Other non-pines
Palms
Unresolved mixed
wetland forests
Unresolved


746


corners


er of
35671
1702
933
142
199
129
468
60
337
85
60
59
1227
756
1604
546
914
284
1451
1578
136
145


1.50


636 1.28


49868


A) ."


% of Total
71.53
3.41
1.87
0.28
0.40
0.26
0.94
0.12
0.68
0.17
0.12
0.12
2.46
1.52
3.22
1.09
1.83
0.57
2.91
3.16
0.27
0.29


24.)


24.1 nresol ed
-------------------------












Table Observed and expected frequencies of community
change between adjacent Land Office Survey sampling points. Plant
communities at adjacent points can either change from pine to
non-pine community, remain pineland community at adjacent points,
or remain non-pineland community at adjacent sampling points.
Expected frequencies were generated by randomly assigning
positions to pine and non-pine communities in the proportion
observed. Expected numbers of each case are based on the mean of
500 simulations. Chi-square test to determine significance.
--------------------------------------------------------------
Type Observed Expected

Pine/Pine 40,925 35,872

Non-Pine/Non-Pine 5,414 3,069

Pine/Non-Pine 13,608 21,006

Chi-square = 5109, df=2, p<.001







/ L












j ?. !





Table 6. A list of species associated with mesic and wetland
hardwood communities.
----------------------------------------------------------------
-----------------------------------------------------------------
# species Association with hardwoods
xeric mesic wetland
N_ ----- ------- -----------------.-_-. .-----
2 Oak xx xx xx
3 Cypress 1 xx!
4 Gum x xx
5 Bay x xx
6 Redbay x 0rx.
7 Sweetbay x 11s6
8 Water Oak 'xx
9 White Oak xx
11 Magnolia xx
12 Hickory xx xx
13 Ash > xx
14 White Bay K xx
15 Beech xx
17 Cedar _xx xx
18 Cherry x xx x
19 Chinquapin xx
21 Dogwood xx x
22 Elm x xx
24 Sweetgum xx x
25 Black Gum x xx
26 Tupelo Gum xx
27 Hackberry xx
28 Haw x xx x
29 Red Haw x xx x
30 Holly xx
31 Hornbeam xx. k
33 Ironwood xx
36 Linden, Linn xx
37 Locust xx
38 Maple x xx
39 Mulberry xx x
40 Myrtle x xx
41 Black Oak xx xx xx
43 Spanish Oak x xx x
44 Swamp Oak x xx
46 Live Oak xx xx xx
47 Willow Oak xx
48 Red Oak xx xx xx
52 Persimmon xx
57 Poplar xx
58 Privet xx
60 Sassafras x xx 7
61 Sourwood xx
62 Whortleberry x .xx ??
64 Willow xx
65 Yaupon x xx
67 Birch xx
75 Titi xx






Table 6. Continued

Species Association with hardwoods
xeric mesic hydric

76 Sumac xx
91 Juniper xx
92 Laurel x xx xx
99 Cucumber Tree xx
107 Baygall xx
-----------------------------------------------------------------































Table 7. General soil types distinguished for north Florida. Data are from
The Environmental Geology Series, Florida Department of Natural Resources,
Bureau of Geology. Accompanying soil type definitions are approximate
proportions of the extent of the study area encompassed by each soil typ
----------------------------------------------------
Soil Type Approximate proportion of corners


1.) Medium Fine Sand 9.5 t

2.) Clayey Sand 49.3 t

3.) Sandy Clay 6.4 %

4.) Shelly Sand and Clay 10.1 %

5.) Gravel and Coarse Sand 11.1 t

6.) Shell Beds 0.2 %

7.) Limestone 9.1 *

8.) Dolomite 0.1

9.) Limestone/Dolomite 4.0 t

10.) Peat 0.1

Authors of map series: Pensacola Sheet; Walter Schmidt
Tallahassee Sheet; Walter Schmidt
Apalachicola Sheet; Walter Schmidt
Valdosta Sheet; Michael S. Knapp
Gainesville Sheet; Michael S. Knapp
Jacksonville Sheet: Thomas M. Scott
Daytona Beach Sheet; Thomas M. Scott




Table 8A. Chi square results for the assoication of the eleven
most abundant presettlement species in north Florida with the
seven major soil types. Counts reported.
----------------------------------------------------------------
Species Soil type
Fine Clayey Sandy Shelly Gravel Lime- Limestone
Sand Sand Clay Sand & & stone & Dolomite
(observed/expected) Clay Sand
----------------------------------------------------------------


Cypress


Gum


Bay


Water Oak


Hickory


Blackjack


Red Oak


(Titi


'I, -.


Pine


Oak


37
85

231
66

98
62


16402 2472
16119 2597


220
266

1054
913

307
407


1423
1437


0
43

13
147

38
66


3468
3770

39
62

171
213

131
95

421
305

74,
64

24
60

291
202


212
275

365
258


517 428
864 139

556 52
362 58


191
116

34
24

4
23

27
77


3386 3207
3162 -3002


55
52

84
180

33
80

126
256

26
54

22
51

371
169


73
50

180
170

98
76

118
243

104
51

54
48

42
161


1204
1475

123
24

194
84

141
37
"t.
169
119

75
25

14
24

16
79


210
284

228
263


Totals 4985 21315 3434 1900

Pearson Chi-square = 4418.7 jef =1\


4181 3970

p < .001






ifPi


a l,


1244 283
1303 210


1951


-----------------------------------------------------------------




Table 8B. Chi square results for the association of the eleven
most abundant presettlement species in north Florida with the
seven major soil types.
-------------------------------------------------------------------
Species Soil type
Fine Clayey Sandy Shelly Gravel Lime- Limestone
Sand Sand Clay Sand & & stone & Dolomite
(observed/expected) Clay Sand
-------------------------------------------------------------------
Pine -- ++ ++ ++
Oak ++ ++ ++
Cypress -- ++ ++ ++ ++
Gum ++ ++ ++ ++
Bay ++ -- ++ ++ -- ++
Water Oak ++ ++ ++ ++
Hickory -- ++ ++
Blackjack ++ -- ++ ++
Red Oak -- ++
Titi ++ -- ++
( ++ ++ -- 00
Totals---------------------------------
Totals J
/" I* 'y





Table 9A. Chi square results for the association of the eleven
most abundant presettlement communities in north Florida with the
seven major soil types. Percent values reported.
----------------------------------------------------------------
Species Soil type
Fine Clayey Sandy Shelly Gravel Lime- Limestone
Sand Sand Clay Sand & & stone & Dolomite
(observed/expected) Clay Sand
----------------------------------------------------------------
Pine 8.70 40.56 6.39 3.47 8.59 8.05 2.75
lands 9.24 40.04 6.31 3.60 7.63 7.70 3.89

Pine/Hick/ .11 1.22 .09 .03 .09 .16 .04
Oak .20 .89 .14 .08 .17 .17 .09

Mixed Hard- .41 1.63 .32 .11 .14 .47 .26
woods .39 1.70 .27 .15 .32 .33 .17

Titi .41 .36 .13 .00 .05 .00 .00
swamps .11 .49 .08 .04 .04 .10 .05

Cypress .23 1.91 .02 .16 .13 .28 .25
swamps .35 1.52 .24 .14 .29 .30 .15

Gum .09 .33 .04 .03 .02 .10 .08
communities .08 .36 .06 .03 .07 .07 .03

Bay .63 1.67 .49 .25 .15 .08 .09
communities .40 1.71 .27 .15 .33 .33 .17

Mixed .78 2.05 .23 .30 .22 .49 .79
Swamps .57 2.48 .39 .22 .47 .48 .24

Palms .00 .00 .00 .00 .01 .06 .45
.06 .27 .04 .02 .05 .05 .03

No Trees .22 .46 .08 .18 .07 .09 .11
.14 .62 .10 .05 .12 .12 .06

Unresolved .17 .81 .25 .05 .25 .14 .14
sites with .21 .92 .15 .08 .17 .18 .09
Pine
---------------------------- -------------------------------------
Totals 11.77 51.00 8.04 4.59 9.72 9.92 4.96

Pearson Chi-square = 5003.1 d.f.=60 p < .001





Table 9B. Chi square results for the association of the eleven
most abundant presettlement communities in north Florida with the
seven major soil types.

Species Soil type
Fine Clayey Sandy Shelly Gravel Lime- Limestone
Sand Sand Clay Sand & & stone & Dolomite
(observed/expected) Clay Sand

1 -- ++ ++ -- ++ ++ -
2 -- ++ -- -- -
3 ++ -- ++ -- ++ ++
4 ++ -- ++ -- ++
5 -- ++ -- ++ -- ++
6 ++ -- 00 -- ++ ++
7 ++ -- ++ ++
8 ++ -- -- ++ -- ++ ++
9 -- -- -- ++ ++
10 ++ -- ++ -- ++
11 ++ ---- ++ -- ++

Totals
































This report is the result of a project supported by the Florida Game and Fresh Water
Fish Commission's Nongame Wildlife Program. Although the report fulfilled the
project's contractual obligations; it has not been reviewed for clarity, style, or
typographical errors, and has not received peer review. Any opinions or
recommendations in this report are those of the authors and do not represent policy of
the Commission.
















Suggested citation:

Schwartz, M.W., J. Travis. 1995. The distribution and character of natural habitats in pre-
settlement northern Florida, as recorded by public land survey records. Fla. Game and
Fresh Water Fish Comm. Nongame Wildl. Program Project Rep. 71 pp + ivi.










(Chapter 2)


March 26, 1990






THE DISTRIBUTION OF NATURAL HABITATS IN PRE-SETTLEMENT NORTHERN
FLORIDA, AS RECORDED BY PUBLIC LAND SURVEY RECORDS.




Mark W. Schwartz* and Joseph Travis





Department of Biological Sciences
Florida State University
Tallahassee, Florida 32306-2043


*-current address:
Department of Botany
Univ. of California, Davis
Davis, CA 95616




for submission to:
Vegetation Science





Key words: Public Land Survey; pre-settlement; longleaf
pine;fire;beech-magnolia climax; Florida





Acknowledgement

We thank the Title and Survey Department of the Florida

Department of Natural Resources for allowing us to use the Public

Land Survey Records, as well as office space. We thank Louise

Robbins, Lori Jones, Debbie Polker, Tom Ostertag, and Bill Fay

for their conscientious work in collating the survey records into

an analyzable form. This study was made possible by grant GFC-86-

020 from the Florida Game and Freshwater Fish Comission, NonGame

Program.




M.W. Schwartz page 4


CHAPTER 2. The original Public Land Survey
record of vegetation in panhandle Florida


INTRODUCTION

The abundance and distribution of plant communities in

panhandle Florida prior to European settlement is. not well

understood. There is general agreement that the coastal plain

vegetation was well represented by longleaf pine communities

(Harper 1914, Gano 1917, Braun 1950, Quarterman & Keever 1962,

Platt et al. 1985, Glitzenstein et al 1987). It has been

established that hardwoods replace longleaf pine forests in the

absence of fire through succession to oak-hickory community, and

then in certain circumstances, to beech-magnolia (Garren 1943,

Braun 1950, Quarterman & Keever 1962, Glitzenstein et al 1987).

However, we lack an understanding of whether the dominance of

longleaf pine was: 1) a response to Native American agriculture

by way of abandoned fields (Quarterman & Keever 1962); 2) a

result of fires set by Native Americans (Braun 1950?); 3) or

primarily a result of natural lightning strike fires (Chapman

1950, Komarek 1964, 1968).

While there is general agreement that the vegetation of

prehistoric northern Florida consisted of a mosaic of longleaf pine

forests interspersed with hardwood forests (Kurz 1942, Braun 1950,

Blaisdell et al, Delcourt and Delcourt 1974, 1977), the actual

abundance of different habitats is disputed. Braun (1950) and

Quarterman and Keever (1962) present beech-magnolia as the climax

vegetation of panhandle Florida. They suggest that beech-magnolia

was potentially widespread. Other studies support this view of a





M.W. Schwartz


potentially widespread hardwood forest using historical data

(Delcourt & Delcourt 1977, Clewell 1986). The opinion that

hardwoods were moderately common in pre-settlement Florida is

reflected in Davis' (1967) map of the natural plant communities

of Florida, despite early accounts to the contrary (Williams

1827, 1833, Cabeza de Vaca, Garcilaso de la Vega, William and

John Bartram, Andre Michaux, Thomas Nuttall, as cited in Small

1921a,b, Harper 1914, 1948, Tebo 1985). Davis characterizes

panhandle Florida as predominantly a mixed hardwood-pineland

forest.

Woodsburning on the coastal plain was frequent during the

early 20th century. Pyne (1982) estimates that 105% of Florida

burned each year during the peak of cattle ranching. These

frequent fires may have changed the abundance and distribution of

hardwoods after settlement. Delcourt and Delcourt (1977),

compared modern and pre-settlement forest composition to find

that beech-magnolia abundance is reduced on the modern landscape.

In addition, widespread commercial planting of pine, has clouded

our ability to resolve the natural plant communities of panhandle

Florida from the modern landscape.

In addition, it is recognized that where fire is excluded in

panhandle Florida, hardwood species dominate (Quarterman and

Keever 1962, Hubbell et al. 1956, Blaisdell et al. 1974).

Widespread fire suppression has been common since the mid-20th

century. Local fire suppression may have allowed hardwood forest

to expand. Pine/oak/hickory forests, common on the modern


page 5




M.W. Schwartz page 6


landscape, may be simply a result of fire suppression.

This study determines the pre-settlement abundance, and "

distribution, of trees and community types in panhandle Florida

through a comprehensive examination of Public Land Survey (PLS)

records. In addition, we examine the patchiness of communities

as well as the association of plant communities with various soil

types.

This reconstruction of pre-settlement vegetation in

panhandle Florida serves as a model for Coastal Plain vegetation

in general. The Spanish restricted settlement to two coastal

communities. As a result, Florida was settled much later than

neighboring Coastal Plain states. Therefore the original survey

methods are more complete, and standardized in Florida than in

most other Coastal Plain states (Bourdo 1956).

DATA and METHODS

Public Land Survey records were collected in panhandle

Florida beginning in 1822, after U.S. acquisition of Florida from

the Spanish. The Spanish had occupied Florida since the late

1500's, however Spanish settlements were restricted to two

coastal communities, St. Augustine and Pensacola (Figure 1).

We define panhandle Florida as that portion of Florida north of

Township 10 south (approximately at the latitude of Gainesville,

Figure 1). We sample all available data within this study region.

Several large portions of data, however, are missing. The largest

of these regions is the Forbes purchase, in the central

panhandle. The Forbes purchase was surveyed independently and

does not include vegetation data. Other missing townships were




M.W. Schwartz page 7



omitted because they were surveyed after local settlement, or

their records are missing.

Townships are 6 miles by 6 miles in size (Figure 2). Each

Township is divided into 36 one square mile sections. At each

section corner four bearing trees were surveyed. Two additional

trees were sampled at the mid-points of each section line. Thus, a

township has 108 points (324 trees) at which vegetation may be

sampled (this includes two of the four exterior township lines as

each exterior line also belongs to an adjacent township). Our

study records the location of all sampled corners and the common

names of trees sampled.

Surveyors marked bearing trees when a survey line was

terminated because of a body of water. We exclude those bearing

trees marked at locations other than section corners and mid-

section points from this analysis. These non-standard data points

bias the PLS sample toward wetland communities owing to the fact

that they were located on the edge of oceans, lakes, streams,

rivers, and swamps.

Our data set consists of 131,227 trees sampled at 49,869

section corners and section mid-points. These section corners

come from over 600 townships. A total of 112 types of bearing

tree designations were encountered, including 7 categories of

non-trees (e.g. mound, pit, no tree, dead tree Table 1).

Surveyors recorded common names of trees. This study uses the set

of common name definitions presented in Table 2. Scientific

names, as well as habitat affinities of species are taken from




M.W. Schwartz page 8

Clewell (1987) and Godfrey (1988). Several of common names are

duplicate for the same species (e.g. bullbay and magnolia both

represent Magnolia grandiflora). An additional 35 species were

sampled fewer than 10 times and are not included in Table 1.

Another suite of uncommon species were simply encoded as "Unusual

Tree" during the data collection process (Table 1). We retain a

list of 65 tree names for this analysis.

Common tree names often lack specificity. For example, the

term 'oak' may refer to any of 25 species. With the exception of

pine, we defer from assigning a species when more than one

species is encompassed by a common name. There are seven species

of pine in Florida (Table 3). Three of the pines frequently grow

in monocultures (longleaf, sand, and pond pine). Sand pine and

pond pine, however are generally restricted to coastal areas and

are relatively unimportant species. In addition, historical

accounts record longleaf pine as the overwhelming dominant pine

of Florida (Small 1921a,b, Harper 1914, 1948).

We use the sample of 2 (mid-section line) or 4 (section

corner) bearing trees to assess the community type at each sample

location. This small sample of trees at each site allows only a

general description of community type, and may result in a

moderate levels of community misclassification. We weight species

by habitat affinity to assign community types to corners. Owing

to the few trees sampled at individual data points we create

several categories of ambiguous assemblages of trees.

Unambiguous common name trees with clear habitat affinities

(e.g. beech, Fagus grandifolia) were assigned a score weighted to




M.W. Schwartz


one of three categories: pineland, mesic hardwoods; or forested

wetland. Ambiguous species names where all species have the same

habitat affinity (e.g. cypress, Taxodium ascendens, T.

distichum) were also assigned a community score. Unambiguous

common name trees with wide habitat affinities (e.g. hackberry,

Celtis laevigata) as well as ambiguous common name trees whose

species range across an array of habitats (e.g. oak, Quercus

sp.), were weighted equally for those habitat types in which it

may be found. A correlation matrix of species in mixed-stand

corners (all single species corners eliminated from correlation

matrix) was used as a check for community assignment. Habitat

affinity scores were totaled for corners, those with clear

habitat affinities were assigned a community type. Those corners

where the assemblage of species could be interpreted as different

habitat types were placed into one of several ambiguous

categories, based on the species sampled. The category of

hardwoods was divided into sub-sets containing the mesic hardwood

species such as beech and magnolia and high hammock species (oaks

and hickories). Wetland communities were also divided in subsets

based on the dominant species at each sample point. Appendix A

contains: 1) species weights for the 69 tree names sampled; 2) a

list of the communities described within the three categories of

pineland, hardwoods, and wetlands; 3) a more complete description

of the classification methods used; and 4) a discussion of

potential misclassifications.

We tested species and communities for association with soil


page 9




M.W. Schwartz page 10

parent material using the Environmental Geology Series (Florida

DNR, Dept. of Geology series). These maps provide a general soil

parent type for each section corner. The soil parent type was

collected for each sample point. We use a contingency table G-

test for non-random association of major species, and

communities, with the common soil types described for panhandle

Florida.

The 49,869 sample locations of bearing trees allows for an

analysis of community distribution. Species and community

distribution figures are plotted using the Tallahassee Base line

as the ordinate, and the Florida Principal Meridian as the

abscissa (Figure 1).

RESULTS

INDIVIDUAL SPECIES

Pine
Table 1 lists abundances of species sampled in the PLS data

from panhandle Florida. Pine, the overwhelming dominant, consists

of 77.0 % (101,360 trees) of the entire sample. Pine was most

often found in association with itself as 71% of the sample

points were exclusively pine. The PLS include 4 unique common

names for pines, of which longleaf was the most frequently

encountered; however, 99.7% of the pines sampled are recorded as

"pine" (Table 1).

Wetland species

The wetland species are a diverse group of moderate abundance

(Table 1). Five tree names (bay, cypress, titi, gum, and water

oak) constitute a total of 10.8% (14,140 trees) of sampled trees





M.W. Schwartz


(Table 1). Of these species Water oak (Quercus nigra, 886 trees),

the cypress' (Taxodium ascendens, and T. distichum, 3801 trees),

and titi's ((Cyrilla racemiflora, C. parviflora, and Cliftonia

monophylla, 1784 trees) are purely wetland species. Bay (6134

trees) may refer any of 4 common species in three genera

(Gordonia lasianthus, Magnolia virginiana, Persea borbonia, P.

palustris). Most species of bay are found exclusively within

wetland communities. Redbay (Persea borbonia) is the exception,

ranging from wetland to mesic sites. Sample points dominated by

bay trees are assumed to be wetlands.

Gum (1535 trees) includes a suite of species in the genus

Nyssa. All species within the genus are wetland species except

for Nyssa sylvatica, which can be found in moist to mesic mixed

hardwood sites. Unlike the other gums, however, N. sylvatica

rarely forms stands, such that corners with only gum are probably

wetlands. Samples with a mixture of trees, of which gum is a

member, may be either mesic mixed hardwoods or a wetland

community.

Four species of ash (Fraxinus caroliniana, F. pauciflora,

F. pennsylvanica, and F. profunda, 570 trees, 0.43% of the

sample) are commonly found in floodplain and river swamps. A

fifth ash, F. americana, is an upland hardwood species. Most ash

sampled were in conjunction with other wetland species. Thus, the

ash are considered the sixth major wetland type.

Upland hardwoods

The most abundant upland species after pine are a suite of


page 11




M.W. Schwartz page 12


oaks representing 6.61% of sampled trees (8672 casds -Table 1).

There are 25 species of oak found in northern Florida (Godfrey

1988). Common names sufficient to define an occurrence to a

single species, or a suite of species common to similar habitats,

appear for 10 species (7708 trees Table 1).

Three of the five most common specific oaks sampled

(blackjack oak -3360 trees, post oak -582 trees, and black oak

-472 trees) are pineland associates. The tree name 'oak' is

applied to an additional 964 trees (Table 1). Many of the oak

occurrences are found exclusively in association with pine. The

oaks found in association exclusively with pine are assumed to

pineland oaks.

The tree name red oak (1511 trees) could apply to four

species, Ouercus falcata. Q_ laevis, O. shumardii, and Q.

velutina. Of these four species, only Q. laevis is common in

pineland communities. The remaining three species are usually

found in mixed hardwood stands. No assumption of community

association is made with red oak. However most red oak

occurrences are in exclusive association with pine and probably

refer to the very abundant Q. laevis.

The hickories (Carva aquatica, C. cordiformis, C. floridana, C.

glabra, C. ovalis, C. pallida, and C. tomentosa) were sampled 860

times for a total of 0.66% of our sample (Table 1). Among the

hickories only C. aquatica is found in floodplain habitats. The

remaining species are found in mesic to dry hardwood forests.

Carya ovalis and C. pallida are rare and local to a small portion

of the study area. Carya cordiformis is a species of more




M.W. Schwartz


northerly distribution and only enters the study area in the

central panhandle, where it is found along river courses. The

majority of the hickories sampled were in association with oaks

and other upland hardwoods, the majority of these hickories are

C. glabra, or C. tomentosa.

The remaining major hardwood species are split between a

diverse group primarily of rich mesic hardwood species. This

group includes dogwood (Cornus florida, C. foemina, 552 trees),

magnolia (Magnolia grandiflora, 441 trees), white oak (Quercus

alba, 391 trees), maple (Acer barbatum, A. leucoderme, A. rubrum,

and A. saccharinum, 354 trees), beech (Faaus grandifolia, 332

trees), holly (Ilex opaca, 316 trees), and sourwood (Oxvdendrum

arboreum, 230 trees). These species are relatively uniform in

abundance. It is interesting to note that beech and magnolia are

infrequently sampled, and not a clear dominant among the mesic

hardwoods of the beech-magnolia climax vegetation. This relative

rarity of beech and magnolia is despite the fact that their

smooth bark would have made them preferred species for blazing as

bearing trees and subject to over-sampling through surveyor bias

(Bourdo 1956).

Among the mesic hardwood tree names, the maples and dogwoods

are each represented by wetland species (A. saccharinum and C.

foemina, respectively). These genera are listed with upland

hardwoods because the majority of PLS records are associated with

upland, rather than wetland species. Box-elder (Acer negundo) is

listed as a separate species and is morphologically different


page 13




M.W. Schwartz


from the other maples such that it is not included under the tree

name of maple.

Palms

Cabbage palm and palmetto were sampled 508 times (0.39 %) in

our PLS data. Palm is primarily associated with .other palms or

pine. Most of these records sample from a pine-palm flatwood

forest, or associated palm-oak hammocks.

FOREST COMMUNITIES

Pineland communities

Pineland samples are separated into four categories; 1) pine

only (2 or 4), 2) three pine with a single individual common to

mesic or dry habitats, 3) a mixture of pine and oak, 4) pineland

oaks (blackjack or post oak, 2 or 4 trees).

Pineland habitats were tested for auto-correlation at the scale

of one-half mile intervals. For this analysis I used the four types

previously classified as pineland (77.5% of 49,868 corners) and the

remaining were classified as non-pineland. I randomly distributed

2,250 (22.5%) non-pineland habitats into a 10,000 cell matrix.

The remaining cells were designated as pineland. I then

calculated the frequency of adjacent cells that sampled: 1) pine

to pine; 2) non-pine to non-pine; and 3) pine to non-pine. I

repeated this simulation 500 times to generate a random

distribution of habitat change frequencies. I compared the null

distribution to the observed frequency with which community type

changed between adjacent corners and found the pineland samples

from panhandle Florida to be significantly clustered (Table 5).

The distribution of pineland communities is presented in


page 14





M.W. Schwartz


Figure 3. Pineland sites are distributed throughout the

panhandle. Pineland communities are less abundant four areas.

First, in the along the major river courses that have wide

floodplains, as we would expect, since floodplain forests contain

very few pines. Second in the Marianna lowlands, where there are

relatively rich soils and calcareous deposits, resulting in

higher frequencies of hardwood forests even today.

The third area where pine is less abundant is in the

Tallahassee red hills region, where extensive Native American

agriculture had changed the landscape considerably by the

beginning of the 19th century (Thebeau 1982, Clewell 1984).

Finally, along the Gulf coast east of the Forbes purchase, an

area rich in palm flatwoods and hardwood swamps.

Oak-Hickory sites

Sites containing some mixture of oak and hickory, or all oak

or hickory were classified into this dry hardwood forest

community type (Table 5). In addition, a category of

pine/oak/hickory was constructed to include corners that sampled

pine and hickory or pine,oak, and hickory. Neither of these

categories was well represented. Oak/hickory was found on 1.5% of

corners and pine/oak/hickory on only .09% of corners. This latter

figure, while very low, does not represent the true amount of

pine/oak/hickory forest. Half of the data points sample only two

trees. None of the mid-section lines could sample a pine, oak,

and a hickory. Further, if pine, oak, and hickory are of equal

abundance and randomly distributed, the probability that a


page 15




M.W. Schwartz page 16


surveyor would sample a pine, an oak, and a hickory while noting

four trees is 0.44. If surveyors were biased in the trees they

selected, or if these species differed in abundance, the

probability could be further reduced that all three species would

be sampled. Our estimate indicates that pine/oak/hickory could

have been as much as 0.2 percent of panhandle Florida. The fact

that oak/hickory, which could sample either a xeric hardwood

community or a successional sere, is rare (1.5%) and pine/hickory

even more rare (.04%) leads us to conclude that the mid-

successional community of pine/oak/hickory was a very

unimportant.

We assign the moderately abundant pine/oak community as a

pineland community. We recognize that some of these sites are

sampled from pine/oak/hickory forest, but given the abundance of

pine/hickory corners the majority should be pinelands.

Pine/oak/hickory is a moderately frequent vegetation type on the

modern landscape. It appears that this abundance is of recent

origin, and not a natural community.

Mesic Mixed Hardwood Communities

A large number of commonly sampled trees fit neatly into the

category of mixed hardwood forest (Table 4). Any combination of

trees at a corner that contain a majority of these species is

classified into this community type.

Mesic communities had the highest number of common species

(>50 occurrences), as well as the largest number of uncommon

species (Table 6). Problematic assemblages within this community

type occur where the species sampled are split between mesic




M.W. Schwartz


hardwood species and species common to other communities. A mixed

pineland-mesic hardwood classification (0.4%) describes some of

these occurrences. These corners probably represent a hardwood

community with a mixture of pines other than longleaf. Those

corners that contain both mesic and wetland hardwoods are

classified into the mixed wetland community (3.2%) along with

corners that sample a mixture of wetland species.

Figure 4 presents the distribution of these community types

across north Florida. Mesic hardwood communities are clustered

within two of the major areas on non-pinelands. First, in the

Marianna redlands region, and also in the Tallahassee red hills

(Figure 1). Other areas of prominent hardwood communities exist

along the Apalachicola River (also observed by Delcourt and

Delcourt 1977), and just west of the Choctawhatchee River in the

western panhandle (Figure 1).

The modern range of beech (Faqus grandifolia) is reported to

extend southward into the study region and terminates with two

outlying populations north of Gainesville (Little 1978). The PLS

data confirm the present range by sampling beech scattered

throughout the northern portion of the study area, and then in

outliers toward peninsular Florida (Figure 5).

Wetlands

The most diverse array of community types include all the

wetland types. Bay (2.9%), gum (0.6%), titi (1.1%), and cypress

(1.8%) swamps are classified separately. Each of these types was

found exclusive of the other major wetland types at many sites.


page 17





M.W. Schwartz


Each of these type was also defined in association with pine.

These pine-wetland associations (2.2%) could represent the

ecotone between wetlands and pineland, or a mixture of lowland

pine within a wetland community. Mixtures of wetland dominants

such as cypress and bay, cypress and gum, and cypress and titi,

were grouped into the mixed wetland types along with sites

sampling a mixture of mesic hardwoods with wetland trees. This

type also included those corners that were dominated by ambiguous

wetland/ mesic hardwoods (e.g. ash or elm). Thus of the 21

communities defined for the purpose of this study 11 are variants

of a wetland community.

Figure 6 presents the locations of all corners that sampled

cypress. Cypress is found along most major waterways shown in

this study, but is most abundant in the eastern portion of the

state along the gulf coast and within the lakes region of the

central highlands. Likewise, cypress dominated communities are

most abundant in the eastern part of the study area (Figure 7).

In contrast, titi is most abundant along the gulf coast west

of the Apalachicola (Figures 8). Sites dominated by gum appear

scattered throughout the study, with no apparent center of

abundance (Figure 9). Sites dominated by bay are also scattered

(Figure 10 & 11). In comparison to the overall distribution of

non-pine sites, bay seems to dominate the northwestern portion of

the study area. Bay is also relatively abundant in the central

panhandle, however there seems to be relatively few bay sites

along the eastern gulf coast, and the central highlands lake

region. Mixed hardwood swamps dominate the Choctawhatchee and


page 18




M.W. Schwartz


Escambia Rivers, and along the eastern gulf coast (Figure 12).

Palms

Palms were almost always found in conjunction with other

palms, pines, or live oak (Table 4). Those corners with palms,

were recorded as separate types (0.4%). Sites with palms are

restricted to limestone and dolomite rich soils along the eastern

gulf coast (Figure 13).

No Trees

Corners with no trees (1.5%, Table 4) sampled indicate one

of two habitat types; beaches or dunes without trees and other

coastal sites, or inland savannas and pitcher plant bogs. Bartram

(1791) reports extensive pitcher plant bogs in panhandle Florida.

The exact locations of these sites are not known. However, the

PLS data samples 761 corners with no trees, a portion of these

may be savanna habitats.

Soils

Both species and communities were tested for non-random

association with soil types as defined by the Environmental

Geology Series (Florida DNR, Div. of Resource Manag., Bureau of

Geology). A total of 10 soil types were defined within the study

area (Table 7). Three of these soil types ( peat, dolomite, and

shell beds) were very rare and eliminated from the chi-square

test of vegetation affinity with soil type. The remaining 7 soil

types were tested against the occurrence of the 11 most abundant

species (Table 8) and the 11 most abundant community types (Table

9). In both cases there is a significant non-random association


page 19




M.W. Schwartz page 20


of vegetation with soils (p<.001, p<.001, Table 8,9). While many

of the oaks and titi show associations with sandiness of the

soil, palms have the strongest association with limestone and

dolomite soils. I eliminated samples where I could not resolve

adjacent soil types.

DISCUSSION

Historical accounts suggest that longleaf pinelands were

very abundant in presettlement north Florida (Williams 1827,

1833, Bartram 1791). Harper (1914) measured pinelands to be the

overwhelming dominant form of vegetation in North Florida in the

early 20th century. Other historical work, using the Public Land

Survey data have indicated otherwise. Delcourt and Delcourt

(1977) used survey data from four townships along the

Apalachicola river to suggest that the hardwood forests were more

abundant in presettlement times than they are today. Likewise,

Clewell (1986) used survey data to collect vegetation information

in the northern Leon County region and found that the forest in

that region was dominated by a Pine/Oak/Hickory mix. He

attributes this to relatively recent abandonment of agriculture

by Miccusukee Indians. The Miccusukee people had extensive

agricultural fields in this region, and had abandoned them after

a raid on their lands in 1704 (Clewell 1986). This area appears

to have been sampled at a time of successional recovery from old

field. Davis (1967) presents a map of presettlement vegetation

for Florida that depicts north Florida as being predominantly a

mixed pine-hardwood region. Our PLS data support the regional

abundance of hardwoods in the study areas of Clewell (1986) and




M.W. Schwartz


Delcourt and Delcourt (1977), but find that results from these

areas previously studied do not apply to the majority of

panhandle Florida. This study supports the anecdotal historical

reports (e.g. Bartram 1791), as well as the less systematic

quantitative measures of community abundance (Williams 1827,

Harper 1914) in measuring pineland as the overwhelming dominant

community-type throughout panhandle Florida.

Those regions with greater abundance of hardwoods are A)

localized to areas of special landforms, such as the Bluffs along

the Apalachicola River (Delcourt and Delcourt 1977.), and the

Marianna lowlands; B) in conjunction with abandoned Native

American agricultural fields; or C) associated with wetlands.

The fact that mid-successional communities such as

pine/oak/hickory are uncommon, and that most hardwood communities

are associated with areas of richer soils or wetlands (A&C above)

suggests that the distribution of hardwood habitats were

relatively stable. This is counter to the hypothesis proposed by

Quarterman and Keever (1962) suggesting that the abundance of

pine was, in part, a result of the abandonment of Native American

agricultural fields. Under this hypothesis we should observe

more habitat in the process of turning over from pineland to

mesic hardwood forest. Further, in the single region where we

have a record of agriculture abandonment pine is under-

represented, not over-represented as this hypothesis predicts.

From a conservation perspective, this study reaffirms that

mesic and wetlands species contain most of the woody dominants of


page 21




M.W. Schwartz page 22


panhandle Florida, and demonstrates that they are found scattered

in low frequencies across the region. Thus populations of this

hardwood diversity is naturally fragmented and patchy in

distribution.






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