Journal of the Oklahoma Native Plant Society, Volume 9, December 2009


Oklahoma Native Plant Record 

Volume 9, December 2009 
38 

Composition and Structure of Bottomland Forest Vegetation at the 
Tiak Research Natural Area, McCurtain County, Oklahoma 

Bruce W. Hoagland
*

Newell Alan McCarty 
Oklahoma Biological Survey and Oklahoma Biological Survey 
Department of Geography  University of Oklahoma 
University of Oklahoma Norman, Oklahoma 73072 
Norman, Oklahoma 73072 
*
Corresponding author 

ABSTRACT 

Although southeastern Oklahoma has substantial areas of bottomland forest, few 
studies exist of this vegetation type. We analyzed forest community structure at the Tiak 
Research Natural Area of the Ouachita National Forest on the Upper Gulf Coastal Plain. 
Vegetation data were collected from 24 12m x 8m macroplots. Percent cover data were 
recorded for ferns, fern-allies, and herbaceous angiosperms. Tree sapling and shrub 
species were recorded in diameter classes, and numbers of stems in estimated height 
classes were recorded for midlevel and canopy trees. One hundred fifty-two taxa of 
woody and herbaceous plants were encountered. Toxicodendron radicans and 
Arundinaria gigantea were the common understory species. Acer rubrum, A. saccharum, 
Carpinus caroliniana, Carya alba, C. texana, Cornus florida, Fraxinus americana, 
Liquidambar styraciflua, and Nyssa sylvatica were common as mid-level species. 
Quercus lyrata and Q. phellos were the most common canopy layer trees. 

INTRODUCTION 

The constituent tree species of 
bottomland forests in Oklahoma varies 
from east to west, as does the upland 
vegetation. Some tree species can be found 
in bottomland forests at most localities 
throughout the state, such as Acer negundo, 
A. saccharum, Celtis laevigata, Fraxinus 
pennsylvanica, and Ulmus americana (Bruner 
1931, Blair and Hubbell 1938, Collins et al. 
1981). But the most diverse assemblages of 
bottomland forest tree species occur in 
eastern Oklahoma, which is fostered by 
numerous oak species (i.e., Quercus lyrata, Q. 
nigra, Q. michauxii, Q. pagoda, Q. palustris, Q. 
phellos, and Q. texana). Quercus lyrata, Q. 
michauxii, and Q. texana are restricted to the 
Upper Gulf Coastal Plain (UGCP) 
extension of southeastern Oklahoma (Blair 
and Hubbell 1938). Other bottomland tree

Hoagland & McCarty 
https://doi.org/10.22488/okstate.17.100068

species limited to this area are Pinus taeda 
and Taxodium distichum (Bruner 1931). 

Despite the diversity of woody and 
understory species in the bottomland forests 
of the UGCP in southeastern Oklahoma, 
few vegetation studies have focused on this 
area. The dominant species in this forest 
type are water tolerant Carya and Quercus 
species, as well as Acer rubrum, Liquidambar 
styraciflua, and Nyssa sylvatica (Bruner 1931; 
Brabander, et al. 1985 ). A quantitative 
analysis of woody vegetation at the Little 
River National Wildlife Refuge (LRNWR), 
located in McCurtain County, identified 
three major forest types: Quercus phellos, 
Carpinus caroliniana, and Taxodium distichum.  
Co-dominant trees at the LRNWR included 
A. rubrum, Liquidambar styraciflua, and Quercus
alba (Hoagland et al. 1996).

The objective of this study was to 
quantify species composition and aspects of 
forest community structure at the TRNA of 



Oklahoma Native Plant Record  
Volume 9, December 2009 

 

  

  Hoagland & McCarty

   

39 

the Ouachita National Forest. The data 
collected are intended to serve as a baseline 
for analysis of temporal change in 
bottomland hardwood forests. This research 
area occupies 80.9 ha in North Caney Creek, 
McCurtain County, Oklahoma (Fig. 1). It 
was established in June of 1990 to protect 
the bottomland hardwood forest 
community and was the first Research 
Natural Area representing the Society of 
American Foresters (SAF) type 88 willow 
oak - water oak - diamondleaf oak cover 
type (Devall 1989). 

The TRNA is located within the UGCP 
physiographic province of southeastern 
Oklahoma (Hunt 1974, Curtis et al. 2008). 
The region is composed of deep alluvial 
deposits underlain by limestones and shales 
of the Washita and Kiamichi formations 
(Davis 1960). Relief ranges from 100.9 to 
102.1 m above sea-level. Soils are composed 
of deep alluvial deposits of Guyton silt loam 
group, a poorly drained soil type found on 
floodplains throughout southeastern 
McCurtain County (Reasoner 1974). The 
climate is Subtropical Humid (Trewartha 
1968) with warm humid summers and a 
mean July temperature of 26.9 °C (80 

o
F) 

and short, mild winters with a mean January 
temperature of 4.0 °C (39 

o
F). Mean annual 

precipitation is 136 cm (53.5 in.; Oklahoma 
Climatological Survey 2009). 

 
METHODS 

 

In order to evaluate habitat structure 
and establish baseline stand conditions, it is 
necessary to collect data listing the species 
present, the number of individuals, and area 
occupied. These types of data are of use to 
ecologists, foresters, and zoologists, as well 
as botanists.  

Prior to field data collection, the 
locations of macroplots were equidistantly 
spaced and established on a 1:24,000 scale 
topographic map. Macroplots were then 
located and established in the field. Plots 
were established in April of 1993 and data 

collected in late April, mid July, and early 
October of the 1993 field season.   

Understory Vegetation data were 
collected from units referred to as 
macroplots, each of which measured 12 m x 
8 m with long axis oriented north to south. 
Microplots, measuring 1.0 m x 1.0 m, were 
placed in each corner of each macroplot. All 
species present in the microplot were 
recorded and percent cover visually 
estimated in 5% increments. The percent 
cover of bryophytes, forest litter, open 
water, and exposed soil was recorded. Cover 
data for ferns, fern-allies, and herbaceous 
angiosperms were also recorded. 

Data for Tree Saplings and Shrub 
species were collected from a subplot within 
the macroplot measuring 8.0 m x 6.0 
m.(26.25 ft. x 19.69 ft). Woody species were 
assigned to the following classes based on 
diameter-at-breast-height (DBH): CL1 = 0 - 
2.0 cm, CL2 = 2.1- 6.0 cm, and CL3 = 6.1 - 
10.0 cm. These data were collected to 
characterize shrub composition and 
regeneration potential of canopy tree 
species. 

Data for Mid-level and Canopy 
species were collected from the entire 
macroplot. The species names of all trees in 
the macroplot were recorded and the height 
of each estimated and assigned to the mid-
level category, defined as trees and shrubs 
3.0 - 15.0m in height; or “canopy”, defined 
as trees in excess of 15.1 m in height. 
Taxonomy follows that of the USDA Plants 
Database (USDA-NRCS 2009). No voucher 
specimens were prepared. 

 
RESULTS AND DISCUSSION 

 

One hundred fifty-two taxa of woody 
and herbaceous plants were encountered at 
the TRNA in the 1993 field season. Thirty-
four were trees or shrubs (22.4%), 12 
woody vines (7.9%), and 106 were 
herbaceous plants (69.7%). There were 134 
(89.3%) perennials, 11 (7.3%) annuals, and 6 
(4.0%) biennials. Carex and Quercus were the 



  Oklahoma Native Plant Record 
Volume 9, December 2009 

Hoagland & McCarty 

40 

largest genera with 10 and 7 species, 
respectively. Galium aparine, Lonicera japonica, 
Morus alba, Vicia villosa, and Trifolium dubium 
were non-native species present at the 
TNRA. The Oklahoma Natural Heritage 
Inventory (2009) tracks nine species at the 
TRNA as state rare, though globally secure: 
Aralia spinosa (G5S1S2), Aristolochia reticulata 
(G4S2), Bignonia capreolata (G5S1), Carex 
debilis (G5S1), Desmodium pauciflorum (G5S1), 
Justicia ovata (G5S?), Quercus texana 
(G4G5S1), Triadenum tubulosum (G4S1S2), 
and Uvularia sessilifolia (G5S1). No federally 
listed threatened or endangered species were 
present. 

 
Understory Vegetation 

Of the 152 species encountered in the 
microplots, 44 (28.9%) were recorded from 
all three sample dates, 50 (32.9%) in two 
seasons, and 58 (38.2%) in one season only 
(Table 1). Of the taxa recorded in one 
season, 23 (39.7%) were in spring only, 
21(36.2%) in summer, and 14 (24.3%) only 
in fall. The highest mean covers were 
“litter” (49.2%) and “exposed soil” 
(16.20%). 

Seven plant taxa scored annual mean 
cover values greater than 1.0%: Toxicodendron 
radicans (5.33%), Arundinaria gigantea (2.23%), 
Vitis vulpina (1.94%), Parthenocissus 
quinquefolia (1.75%), Quercus phellos seedlings 
(1.63%), Carex sp. (1.20%), and Acer rubrum 
(1.15%). A total of 11 Carex species were 
identified, with mean cover values ranging 
from 0.55% (C. gravida) to 0.01% (C. 
laxiflora). 

Mean cover values for each sample 
period were comparable; spring, 41%; 
summer, 38%; fall, 33%. There was greater 
variance in the number of species 
encountered between sampling periods. The 
most species were recorded for the summer 
(108), followed closely by the spring (101), 
and fall (83). Toxicodendron radicans had the 
highest mean cover for all three sampling 
periods. In the spring, T. radicans (6.94%), 
Parthenocissus quinquefolia (3.5%), and Vitis 

vulpina (3.46%) had the largest average 
cover. Likewise in the summer, Toxicodendron 
radicans (7.58%) had a substantially larger 
cover average value than Arundinaria gigantea 
(3.82%) and Quercus phellos seedlings 
(2.43%). Fewer species were recorded in the 
fall. Only in the fall data did other species 
outscore T. radicans in average cover: A. 
gigantea (2.01%), Carex sp. (1.74%), Q. phellos 
(1.65%) and Q. nigra (1.50%) seedling T.  
radicans (1.47%), and Chasmanthium latifolium 
(1.46%). Nevertheless, litter and bare 
ground had the highest percent cover in all 
seasons. 

 
Tree Sapling and Shrub 

Nineteen woody species and 281 stems 
were recorded in the shrub/sapling plots 
(Table 2). All 19 species were represented in 
CL1, but only 13 in CL2 and 10 in CL3. 
Nine species occurred in all three DBH 
classes; Acer rubrum, A. saccharum, Carpinus 
caroliniana, Carya alba, C. texana, Cornus 
florida, Fraxinus americana, Liquidambar 
styraciflua, and Nyssa sylvatica. With the 
exception of C. texana, these species are 
typical of bottomland forest habitats on the 
UGCP. 

Each of the three species with 20 or 
more stems represents one level in the 
forest: Rhododendron canescens, shrub; Ostrya 
virginiana, mid-level; and Nyssa sylvatica, 
canopy. No tree species exceeded 20 stems 
in the remaining DBH classes.  Liquidambar 
styraciflua, a species of second growth 
bottomland forests, had the greatest number 
of stems in CL2 and Carpinus caroliniana, a 
common understory tree of bottomland 
forests, in CL3. 
 
Mid-level and Canopy 

Twenty-three species and 3,797 stems 
were included in the two height categories 
(Table 3). Twelve species occurred in both 
the mid-level and canopy categories. 
Twenty-two species and the majority of 
stems (2,087; 55%) were in the mid-level 
category. Liquidambar styraciflua had the most 



Oklahoma Native Plant Record  
Volume 9, December 2009 

 

  

  Hoagland & McCarty

   

41 

stems (385; 18.4%). The stem counts for 
Acer rubrum (275, 13.2%), C. texana (245, 
11.7%), Q. alba (220, 10.5%), and N. sylvatica 
(210, 10.1%) were also high, but 
substantially less than L. styraciflua.  

The canopy category consisted of 12 
species and 1,710 (45%) stems. Quercus 
texana was the only species in the canopy 
category that was not in the mid-level 
category. Quercus lyrata (330 stems, 19.3%) 
and Q. phellos (315 stems, 18.4%) were the 
most abundant species in the canopy layer. 
Quercus rubra (265 stems, 15.5%) and 
Liquidambar styraciflua (260 stems, 15.2%) 
were also common in the canopy of TRNA. 
Acer rubrum, represented by 275 stems in the 
mid-level, had only 30 stems (1.8%) in the 
canopy. Of the most prominent trees in the 
mid-level, only Quercus alba had similar 
numbers of stems in the canopy (220 vs. 
195). 

 
CONCLUSIONS 

 

Fewer woody plant species were found 
at the TRNA (27) than the Little River 
National Wildlife Refuge (LRNWR; 47; 
Hoagland et al. 1996). Furthermore, with 
the exception of Asimina triloba and 
Rhododendron canescens, all woody species 
reported from TRNA were present at the 
LRWNR. The disparity in species numbers 
is intriguing. Although the LRNWR is larger 
than the TRNA, it was sampled with fewer 
plots. The greater number of species at 
LRNWR might reflect a higher degree of 
habitat variability. The TRNA is 
predominately inundated to seasonally 
inundated habitat, but also includes upland 
vegetation, though limited to the southeast 
(plots 4, 5, 8, and 9) and northeast corners 
of the site (plots 19, 20, 21, and 22; see Fig. 
1). 

The macroplot approach adopted for 
this study was successful for assessing the 
predominant plant species present at the 
TRNA, and data collected from microplots 
for herbaceous species were informative. 

However there are shortcomings in the 
types of data collected for woody plant 
species. Rather than assigning species to 
pre-established classes, actual measurement 
of tree and shrub DBH and height would 
allow for thorough analysis of woody 
species composition and forest structure. 
Likewise, a detailed inventory of vascular 
plants in the TRNA would benefit future 
monitoring projects. 
 

LITERATURE CITED 
 

Blair WF and Hubbell TH. 1938. The biotic 
districts of Oklahoma. American Midland 
Naturalist 20:425-454. 

Brabander JJ, Masters RE, and Short RM. 
1985. Bottomland hardwoods of eastern 
Oklahoma. U. S. Fish Wildlife Service, 
Tulsa, OK. 145 pp. 

Bruner WE. 1931. The vegetation of 
Oklahoma. Ecological Monographs 1:99-
188. 

Collins SL, Risser PG, and Rice EL. 1981. 
Ordination and classification of mature 
bottomland forests in north central 
Oklahoma. Bulletin of the  Torrey Botanical 
Club 108:152-165. 

Curtis NM, Ham, WE, and Johnson KS. 
2008. Geomorphic provinces of 
Oklahoma. In: Johnson, K.S. and K.V. 
Luza (eds.). Earth sciences and mineral 
resources of Oklahoma. Oklahoma 
Geological Survey, Norman, Oklahoma. 

Davis, L. 1960. Geology and groundwater 
resources of southern McCurtain 
County, Oklahoma. Oklahoma Geological 
Survey. Bulletin 86. 

Devall, MS 1989. Establishment record for the 
Tiak Research Natural Area within the 
Ouachita National Forest, McCurtain 
County, Oklahoma. Southern Forest 
Experiment Station, U. S. Forest 
Service, New Orleans. 

Hoagland BW, Sorrels LR, and Glenn SM. 
1996. Woody species composition of 
floodplain forests of the Little River, 
McCurtain and LeFlore counties, 



  Oklahoma Native Plant Record 
Volume 9, December 2009 

Hoagland & McCarty 

42 

Oklahoma. Proceedings of the Oklahoma 
Academy of Science 76:23-26. 

Hoagland BW, Buthod A, Callahan-
Crawford P, Elisens E, and Tyrl R.  
2009. Oklahoma vascular plants database. 
(www.biosurvey.ou.edu) University of 
Oklahoma, Norman. (Accessed 1 
January 2009). 

Hunt, CB. 1974. Natural regions of the United 
States and Canada. W.H. Freeman, San 
Francisco. 725 p. 

Oklahoma Climatological Survey. 2009. 
Oklahoma Climatological Data. 
(www.ocs.ou.edu) University of 
Oklahoma, Norman. (Accessed 1 
November 2009). 

Oklahoma Natural Heritage Inventory 
(ONHI). 2009. Oklahoma Natural 
Heritage Inventory working list of rare 
Oklahoma plants. 
(www.biosurvey.ou.edu/publicat.html) 
University of Oklahoma, Norman. 
(Accessed 1 November 2009). 

Reasoner, RC 1974. Soil survey of McCurtain 
County, Oklahoma. United States 
Department of Agriculture, 
Washington, D.C. 

USDA-NRCS. 2008. The plants database. 
(www.plants.usda.gov) National Plant 
Data Center, Baton Rouge, LA. 

Trewartha, GT 1968. An introduction to 
climate. McGraw-Hill, New York.  

 

 

 
 
Figure  Location of the Tiak Research Natural Area, Ouachita National Forest, McCurtain 
County, Oklahoma. Numbers indicate locations of macroplots. 

 

www.biosurvey.ou.edu
www.ocs.ou.edu
http://www.biosurvey.ou.edu/publicat.html
http://www.plants.usda.gov/


 

 

43 

 

Table 1  Species composition of microplots at the Tiak  Research Natural Area, Ouachita National Forest, McCurtain County, Oklahoma. Columns denoted 
represent average cover value for all seasons (MEAN), number of macroplots (n=24) in which a species was encountered (FREQ), and percentage or relative 
frequency (RF) of macroplots in which a species occurred. The remaining columns provided the same data for the spring (April), summer (July), and fall (October) 
sampling periods. 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Acalypha rhomboidea 0.04 2  8.3 
 

0.02 1 4 
 

0.01 1 4 
 

0.10 1 4 

Acer rubrum 1.15 19 79.2 
 

1.42 14 58 
 

1.29 12 50 
 

0.74 14 58 

Acer saccharum 0.24 7 29.2 
 

0.04 1 4 
 

0.38 5 21 
 

0.30 6 25 

Agrimonia parviflora 0.09 2  8.3 
 

0.23 2 8 
 

0.04 1 4 
 

0.00 0 0 

Agrostis hyemalis 0.08 3 12.5 
 

0.00 0 0 
 

0.13 2 8 
 

0.10 3 13 

Allium canadense 0.14 5 20.8 
 

0.41 6 25 
 

0.00 0 0 
 

0.00 0 0 

Aralia spinosa 0.01 1  0.0 
 

0.00 0 0 
 

0.04 1 4 
 

0.00 0 0 

Arisaema dracontium  0.01 2  8.3 
 

0.04 2 8 
 

0.00 0 0 
 

0.00 0 0 

Aristida sp. 0.23 1  4.2 
 

0.00 0 0 
 

0.68 2 8 
 

0.00 0 0 

Aristolochia reticulata 0.04 2  8.3 
 

0.00 0 0 
 

0.05 1 4 
 

0.06 2 8 

Arundinaria gigiantea 2.23 11 45.8 
 

0.84 9 38 
 

3.82 8 33 
 

2.01 10 42 

Asimina triloba 0.20 4 16.7 
 

0.17 3 13 
 

0.13 3 13 
 

0.30 4 17 

Athyrium filix-femina 0.01 1  4.2 
 

0.04 1 4 
 

0.00 0 0 
 

0.00 0 0 



44 

 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Berchemia scandens  0.56 11 45.8 
 

0.56 8 33 
 

0.54 7 29 
 

0.58 10 42 

Betula nigra 0.03 1  4.2 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 1 4 

Bignonia capreolata 0.28 5 20.8 
 

0.00 0 0 
 

0.26 3 13 
 

0.59 5 21 

Boehmeria cylindrica 0.30 4 16.7 
 

0.00 0 0 
 

0.78 5 21 
 

0.12 4 17 

Botrychium virginianum 0.11 5 20.8 
 

0.05 1 4 
 

0.00 0 0 
 

0.27 7 29 

Callicarpa americana 0.19 6 25.0 
 

0.04 1 4 
 

0.25 6 25 
 

0.27 7 29 

Campsis radicans 0.39 8 33.3 
 

0.00 0 0 
 

0.61 5 21 
 

0.56 8 33 

Carex complanata 0.55 15 62.5 
 

1.54 14 58 
 

0.00 0 0 
 

0.10 1 4 

Carex debilis 0.36 6 25.0 
 

1.07 1 4 
 

0.00 0 0 
 

0.00 0 0 

Carex gravida 0.65 18 75.0 
 

0.81 11 46 
 

0.51 10 42 
 

0.63 12 50 

Carex grayi 0.07 1  4.2 
 

0.00 0 0 
 

0.22 1 4 
 

0.00 0 0 

Carex intumescens 0.38 11 45.8 
 

0.74 8 33 
 

0.00 0 0 
 

0.40 7 29 

Carex laxiflora 0.01 0  0.0 
 

0.00 0 0 
 

0.02 1 4 
 

0.00 0 0 

Carex lupulina 0.54 15 62.5 
 

0.10 2 8 
 

1.35 13 54 
 

0.18 2 8 

Carex oligocarpa 0.18 9 37.5 
 

0.00 0 0 
 

0.43 8 33 
 

0.10 1 4 



 

 

45 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Carex sp. 1.20 21 87.5 
 

1.67 1 4 
 

0.19 2 8 
 

1.74 14 58 

Carex squarrosa 0.03 3 12.5 
 

0.04 1 4 
 

0.06 2 8 
 

0.00 0 0 

Carex tribuloides 0.28 5 20.8 
 

0.00 0 0 
 

0.83 1 4 
 

0.00 0 0 

Carpinus caroliniana 0.39 10 41.7 
 

0.42 5 21 
 

0.38 6 25 
 

0.39 4 17 

Carya alba 0.05 4 16.7 
 

0.02 1 4 
 

0.13 3 13 
 

0.00 0 0 

Carya aquatica 0.05 3 12.5 
 

0.11 2 8 
 

0.04 1 4 
 

0.00 0 0 

Carya sp. 0.19 9 37.5 
 

0.15 5 21 
 

0.00 0 0 
 

0.42 10 42 

Carya texana 0.26 12 50.0 
 

0.23 3 13 
 

0.45 12 50 
 

0.10 1 4 

Cercis canadensis 0.05 1  4.2 
 

0.06 1 4 
 

0.00 0 0 
 

0.10 1 4 

Chasmanthium latifolium 0.05 2  8.3 
 

0.00 1 4 
 

0.15 2 8 
 

0.00 0 0 

Chasmanthium laxum 0.40 8 33.3 
 

0.05 1 4 
 

1.14 9 38 
 

0.00 0 0 

Chasmanthium 
sessiliflorum 0.53 10 41.7 

 
0.00 0 0 

 
0.12 2 8 

 
1.46 11 46 

Clitoria mariana 0.04 2  8.3 
 

0.00 0 0 
 

0.05 2 8 
 

0.08 2 8 

Convolvulus sp. 0.02 1  4.2 
 

0.00 0 0 
 

0.06 2 8 
 

0.00 0 0 

Cornus florida 0.15 7 29.2 
 

0.24 6 25 
 

0.21 3 13 
 

0.00 0 0 



46 

 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Crataegus marshallii 0.17 2  8.3 
 

0.26 3 13 
 

0.12 3 13 
 

0.14 3 13 

Desmodium nudiflorum 0.07 2  8.3 
 

0.00 0 0 
 

0.10 2 8 
 

0.10 1 4 

Desmodium pauciflorum 0.03 1  4.2 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 2 8 

Desmodium sp. 0.08 4 16.7 
 

0.00 0 0 
 

0.13 4 17 
 

0.10 1 4 

Dichanthelium 
acuminatum var. 
fasciculatum 0.01 2  8.3 

 

0.00 0 0 

 

0.03 2 8 

 

0.00 0 0 

Dichanthelium boscii 0.18 12 50.0 
 

0.03 1 4 
 

0.52 12 50 
 

0.00 0 0 

Dioscorea quaternata 0.68 16 66.7 
 

0.95 10 42 
 

0.98 14 58 
 

0.10 1 4 

Diospyros virginiana 0.07 3 12.5 
 

0.11 2 8 
 

0.00 0 0 
 

0.08 2 8 

Elephantopus 
carolinianus 0.12 6 25.0 

 
0.00 0 0 

 
0.23 5 21 

 
0.13 5 21 

Elephantopus 
tomentosus 0.01 2  8.3 

 
0.00 0 0 

 
0.04 1 4 

 
0.00 0 0 

Elymus virginicus 0.02 0  0.0 
 

0.05 1 4 
 

0.00 0 0 
 

0.00 0 0 

Euonymus americana 0.16 7 29.2 
 

0.07 2 8 
 

0.27 6 25 
 

0.13 4 17 

Fraxinus americana 0.07 3 12.5 
 

0.13 1 4 
 

0.04 1 4 
 

0.05 1 4 

Galium aparine 0.07 5 20.8 
 

0.13 2 8 
 

0.08 3 13 
 

0.00 0 0 



 

 

47 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Galium circaezans 0.08 3 12.5 
 

0.24 4 17 
 

0.00 0 0 
 

0.00 0 0 

Galium concinnum 0.14 6 25.0 
 

0.42 6 25 
 

0.00 0 0 
 

0.00 0 0 

Geum canadense 0.15 6 25.0 
 

0.26 3 13 
 

0.10 3 13 
 

0.10 2 8 

Gillenia stipulata 0.01 1  4.2 
 

0.00 0 0 
 

0.02 1 4 
 

0.00 0 0 

Gratiola neglecta  0.03 2  8.3 
 

0.08 2 8 
 

0.00 0 0 
 

0.00 0 0 

Hypericum hypericoides 0.17 6 25.0 
 

0.05 1 4 
 

0.15 5 21 
 

0.29 7 29 

Hypericum mutilum 0.03 0  0.0 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 0 0 

Hypericum punctatum 0.03 1  4.2 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 1 4 

Ilex opaca 0.14 4 16.7 
 

0.11 3 13 
 

0.17 3 13 
 

0.14 4 17 

Impatiens capensis 0.50 13 54.2 
 

1.49 13 54 
 

0.00 0 0 
 

0.00 0 0 

Juncus coriaceus 0.01 1  4.2 
 

0.04 1 4 
 

0.00 0 0 
 

0.00 0 0 

Justicia ovata 0.64 5 20.8 
 

1.62 1 4 
 

0.20 4 17 
 

0.09 2 8 

Lactuca canadensis 0.02 2  8.3 
 

0.03 2 8 
 

0.04 1 4 
 

0.00 0 0 

Lactuca floridana 0.03 3 12.5 
 

0.06 2 8 
 

0.02 1 4 
 

0.00 0 0 

Lactuca sp. 0.01 1  4.2 
 

0.00 0 0 
 

0.04 1 4 
 

0.00 0 0 



48 

 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Liquidambar styraciflua 0.43 15 62.5 
 

0.30 9 38 
 

0.42 11 46 
 

0.56 13 54 

Lonicera japonica 0.03 2  8.3 
 

0.00 0 0 
 

0.00 0 0 
 

0.08 3 13 

Maianthemum 
racemosum 0.03 1  4.2 

 
0.06 1 4 

 
0.02 1 4 

 
0.00 0 0 

Matelea biflora 0.08 3 12.5 
 

0.23 3 13 
 

0.00 0 0 
 

0.00 0 0 

Mitchella repens 0.23 7 29.2 
 

0.11 3 13 
 

0.15 2 8 
 

0.44 6 25 

Monarda punctata 0.02 1  4.2 
 

0.00 0 0 
 

0.00 0 0 
 

0.05 1 4 

Monarda sp. 0.03 1  4.2 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 1 4 

Monarda russeliana 0.01 0  0.0 
 

0.00 0 0 
 

0.04 1 4 
 

0.00 0 0 

Morus alba 0.05 2  8.3 
 

0.04 1 4 
 

0.02 1 4 
 

0.10 1 4 

Morus rubra 0.03 1  4.2 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 1 4 

Muhlenbergia sp. 0.03 2  8.3 
 

0.00 0 0 
 

0.08 2 8 
 

0.00 0 0 

Nyssa sylvatica 0.12 6 25.0 
 

0.00 0 0 
 

0.31 5 21 
 

0.05 2 8 

Oligoneuron nitidum 0.19 0  0.0 
 

0.09 2 8 
 

0.26 1 4 
 

0.20 2 8 

Onoclea sensibilis 0.02 2  8.3 
 

0.04 1 4 
 

0.01 1 4 
 

0.00 0 0 

Ostrya virginiana 0.42 8 33.3 
 

0.50 7 29 
 

0.66 5 21 
 

0.10 1 4 



 

 

49 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Oxalis stricta 0.13 9 37.5 
 

0.40 10 42 
 

0.00 0 0 
 

0.00 0 0 

Packera obovata 0.02 2  8.3 
 

0.02 1 4 
 

0.04 1 4 
 

0.00 0 0 

Panium anceps 0.25 12 50.0 
 

0.56 10 42 
 

0.00 0 0 
 

0.19 5 21 

Parthenocissus 
quinquefolia 1.75 16 66.7 

 
3.50 15 63 

 
1.64 12 50 

 
0.10 1 4 

Passiflora lutea 0.09 7 29.2 
 

0.00 0 0 
 

0.18 6 25 
 

0.10 2 8 

Penstemon digitalis 0.09 1  4.2 
 

0.24 3 13 
 

0.04 1 4 
 

0.00 0 0 

Penthorum sedoides 0.02 1  4.2 
 

0.00 0 0 
 

0.05 1 4 
 

0.00 0 0 

Poa autumnalis 0.14 9 37.5 
 

0.31 7 29 
 

0.11 2 8 
 

0.00 0 0 

Podophyllum peltatum 0.01 1  4.2 
 

0.02 1 4 
 

0.00 0 0 
 

0.00 0 0 

Polygonatum biflorum 0.01 1  4.2 
 

0.00 0 0 
 

0.01 1 4 
 

0.00 0 0 

Polygonum 
hydropiperoides 0.01 1  4.2 

 
0.00 0 0 

 
0.02 1 4 

 
0.00 0 0 

Polygonum virginianum 0.17 5 20.8 
 

0.11 2 8 
 

0.18 5 21 
 

0.21 6 25 

Prunella vulgaris 0.01 1  4.2 
 

0.00 0 0 
 

0.03 1 4 
 

0.00 0 0 

Prunus serotina 0.05 2  8.3 
 

0.06 2 8 
 

0.04 1 4 
 

0.05 2 8 

Pycnanthemum 
tenufolium 0.04 1  4.2 

 
0.08 2 8 

 
0.04 1 4 

 
0.00 0 0 



50 

 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Quercus alba 0.23 10 41.7 
 

0.20 6 25 
 

0.40 9 38 
 

0.10 2 8 

Quercus lyrata 0.05 4 16.7 
 

0.00 0 0 
 

0.10 5 21 
 

0.06 1 4 

Quercus nigra 0.92 17 70.8 
 

0.32 7 29 
 

0.95 13 54 
 

1.50 13 54 

Quercus phellos 1.63 20 83.3 
 

0.83 18 75 
 

2.43 13 54 
 

1.65 10 42 

Quercus rubra 0.02 3 12.5 
 

0.05 3 13 
 

0.01 1 4 
 

0.00 0 0 

Quercus sp. 0.33 16 66.7 
 

0.36 11 4 
 

0.00 0 0 
 

0.62 11 46 

Quercus velutina 0.11 1  4.2 
 

0.33 1 4 
 

0.00 0 0 
 

0.00 0 0 

Ranunculus abortivus 0.03 3 12.5 
 

0.10 3 13 
 

0.00 0 0 
 

0.00 0 0 

Rhododendron 
canescens 0.24 6 25.0 

 
0.40 6 25 

 
0.20 3 13 

 
0.14 2 8 

Rosa sp. 0.05 3 12.5 
 

0.03 1 4 
 

0.05 2 8 
 

0.06 3 13 

Rubus trivalis 0.29 9 37.5 
 

0.38 6 25 
 

0.20 2 8 
 

0.30 6 25 

Salvia lyrata 0.03 3 12.5 
 

0.06 3 13 
 

0.03 2 8 
 

0.00 0 0 

Sanicula canadensis 0.04 4 16.7 
 

0.00 0 0 
 

0.13 5 21 
 

0.00 0 0 

Sassifras albidum 0.03 1  4.2 
 

0.07 1 4 
 

0.02 1 4 
 

0.00 0 0 

Saururus cernuus 0.33 3 12.5 
 

0.25 2 8 
 

0.47 3 13 
 

0.26 1 4 



 

 

51 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Scleria oligantha 0.18 8 33.3 
 

0.00 0 0 
 

0.55 9 38 
 

0.00 0 0 

Sisyrinchium 
angustifolium 0.01 1  4.2 

 
0.03 1 4 

 
0.00 0 0 

 
0.00 0 0 

Smilax bona-nox 0.24 14 58.3 
 

0.34 10 42 
 

0.37 9 38 
 

0.00 0 0 

Smilax glauca 0.32 16 66.7 
 

0.05 1 4 
 

0.15 3 13 
 

0.76 17 71 

Smilax rotundifolia 0.48 18 75.0 
 

0.35 11 46 
 

0.70 15 63 
 

0.39 9 38 

Smilax sp.  0.05 4 16.7 
 

0.00 0 0 
 

0.06 3 13 
 

0.10 1 4 

Solidago sp. 0.09 2  8.3 
 

0.00 0 0 
 

0.00 5 21 
 

0.28 3 13 

Symphyotrichum 
cordifolium 0.02 1  4.2 

 
0.00 0 0 

 
0.05 1 4 

 
0.00 0 0 

Symphyotrichum 
praealtum 0.08 3 12.5 

 
0.05 1 4 

 
0.18 4 17 

 
0.02 1 4 

Symphyotrichum pratens 0.01 0  0.0 
 

0.00 0 0 
 

0.04 1 4 
 

0.00 0 0 

Symphyotrichum sp. 0.18 4 16.7 
 

0.10 1 4 
 

0.02 1 4 
 

0.41  4 17 

Thalictrum thalictroides 0.02 1  4.2 
 

0.03 1 4 
 

0.04 1 4 
 

0.00 0 0 

Tilia americana 0.17 4 16.7 
 

0.31 1 4 
 

0.10 3 13 
 

0.50 1 4 

Toxicodendron radicans 5.33 21 87.5 
 

6.94 19 79 
 

7.58 18 75 
 

1.47 16 67 

Tradescantia ohiensis 0.10 2  8.3 
 

0.23 3 13 
 

0.06 2 8 
 

0.00 0 0 



52 

 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Triadenum tubulosum 0.19 2  8.3 
 

0.00 0 0 
 

0.35 3 13 
 

0.22 2 8 

Trifolium dubium 0.03 0  0.0 
 

0.00 0 0 
 

0.00 0 0 
 

0.10 1 4 

Ulmus alata 0.03 2  8.3 
 

0.00 0 0 
 

0.00 0 0 
 

0.08 2 8 

Ulmus rubra 0.18 10 41.7 
 

0.00 0 0 
 

0.00 0 0 
 

0.54 12 50 

Ulmus serotina 0.11 5 20.8 
 

0.10 1 4 
 

0.21 6 25 
 

0.00 0 0 

Uvularia sessiliflora 0.01 1  4.2 
 

0.04 1 4 
 

0.00 0 0 
 

0.00 0 0 

Vaccinium corymbosum 0.05 2  8.3 
 

0.04 1 4 
 

0.10 1 4 
 

0.00 0 0 

Vaccinium stamineum 0.60 3 12.5 
 

0.21 3 13 
 

0.94 3 13 
 

0.65 4 17 

Viburnum rufidulum 0.02 1  4.2 
 

0.06 1 4 
 

0.00 0 0 
 

0.00 0 0 

Vicia villosa 0.02 2  8.3 
 

0.05 2 8 
 

0.00 1 4 
 

0.00 1 4 

Viola bicolor 0.33 13 54.2 
 

0.99 13 54 
 

0.00 0 0 
 

0.00 0 0 

Viola pubescens 0.01 1  4.2 
 

0.02 1 4 
 

0.00 0 0 
 

0.00 0 0 

Viola sp. 0.05 3 12.5 
 

0.05 1 4 
 

0.11 4 17 
 

0.00 0 0 

Vitis aestivalis 0.12 2  8.3 
 

0.03 1 4 
 

0.25 2 8 
 

0.08 1 4 

Vitis vulpina 1.94 19 79.2 
 

3.46 13 54 
 

1.02 13 54 
 

1.34 18 75 



 

 

53 

 

 MEAN 

 

ANNUAL 

 

FREQ RF 

 

MEAN 

 

SPRING 

 

FREQ RF 

 

MEAN 

 

SUMMER 

 

FREQ RF 

 

MEAN 

 

FALL 

 

FREQ RF 

Woodwardia areolata 0.13 2  8.3 
 

0.00 0 0 
 

0.33 2 8 
 

0.05 1 4 

Zizea aurea 0.08 4 16.7 
 

0.23 4 17 
 

0.00 0 0 
 

0.00 0 0 

Bryophytes  4.61 8 33.3 
 

 3.87 5 21 
 

 5.95 8 33 
 

 4.01 7 29 

Exposed soil 16.29 20 83.3 
 

18.19 18 75 
 

20.02 20 83 
 

10.65 20 83 

Organic litter 49.24 19 79.2 
 

18.19 19 79 
 

57.57 22 92 
 

71.98 19 79 

Pooled water  1.90 4 16.7 
 

 2.53 3 13 
 

 0.04 1 4 
 

 3.13 1 4 

 



54 

 

 

Table 2  Tree sapling and shrub species composition at the Tiak Research Natural Area, Ouachita National Forest, McCurtain County, Oklahoma.  
Columns denoted number of stems within each DBH class (CL1 = 0-2.0 cm, CL2 = 2.1-6.0 cm, and CL3 = 6.1-10.0 cm), number of macroplots (n=24)  
in which a species was encountered (FREQ), and percentage or relative frequency (RF)of macroplots in which a species occurred. 

 

 

 

 

#Stems 

 

CL1 

 

FREQ RF 

 

#Stems 

CL2 

 

FREQ RF 

 

#Stems 

CL3 

 

FREQ RF 

Acer rubrum 15 11 45.8 

 

3 2  8.3 

 

5 4 16.7 

Acer saccharum 16 5 20.8 

 

3 1  4.2 

 

2 2  8.3 

Asimina triloba 4 1  4.2 

 

0 0  0.0 

 

0 0  0.0 

Carpinus caroliniana 19 8 33.3 

 

7 7 29.2 

 

8 7 29.2 

Carya alba 11 6 25.0 

 

7 6 25.0 

 

6 5 20.8 

Carya aquatica 1 1  4.2 

 

0 0  0.0 

 

0 0  0.0 

Carya texana 12 9 37.5 

 

9 6 25.0 

 

2 2  8.3 

Cersis canadensis 3 1  4.2 

 

0 0  0.0 

 

0 0  0.0 

Cornus florida 6 3 12.5 

 

1 1  4.2 

 

1 1  4.2 

Euonymus americana 1 1  4.2 

 

0 0  0.0 

 

0 0  0.0 



 

 

55 

 

 

 

 

 

#Stems 

 

CL1 

 

FREQ RF 

 

#Stems 

CL2 

 

FREQ RF 

 

#Stems 

CL3 

 

FREQ RF 

Fraxinus americana 2 2  8.3 

 

2 2  8.3 

 

1 1  4.2 

Ilex opaca 8 5 20.8 

 

1 1  4.2 

 

0 0  0.0 

Liquidambar styraciflua 7 6 25.0 

 

10 5 20.8 

 

6 5 20.8 

Nyssa sylvatica 25 12 50.0 

 

8 5 20.8 

 

5 4 16.7 

Ostrya virginiana 21 9 37.5 

 

1 1  4.2 

 

0 0  0.0 

Quercus alba 3 3 12.5 

 

0 0  0.0 

 

1 1  4.2 

Quercus nigra 1 1  4.2 

 

0 0  0.0 

 

0 0  0.0 

Rhododendron canescens 21 4 16.7 

 

1 1  4.2 

 

0 0  0.0 

Ulmus alata 7 4 16.7 

 

7 4 16.7 

 

0 0  0.0 

Total stems 184   

 

60   

 

37   



56 

 

 

Table 3  Woody species composition within Mid-level and Canopy height classes at the Tiak  Research Natural Area, Ouachita National Forest, McCurtain  
County, Oklahoma. Mid-level woody plants are defined as trees and shrubs 3-15 m in height and Canopy as trees in excess of 16 m. The first column in each 
category represents the number of macroplots (n=24) in which a species was encountered (FREQ), followed by the percentage or relative frequency (RF) of 
macroplots in which a species occurred, the total number of stems (#Stems) and mean number of stems (MStems) for a species. 

 

 

Mid-level 

 

FREQ RF #Stems MStems 

 

Canopy 

 

FREQ RF #Stems MStems 

Acer rubrum 10 42 275 11.4 

 

0 0 30  1.2 

Acer saccharum 3 13 55  2.3 

 

0 0 0  0.0 

Carpinus caroliniana 4 17 100  4.2 

 

0 0 0  0.0 

Carya alba 4 17 105  4.4 

 

0 0 0  0.0 

Carya aquatica 1 4 35  1.5 

 

1 4 40  1.7 

Carya texana 11 46 245 10.2 

 

1 4 25  1.0 

Cornus florida 3 13 15  0.6 

 

0 0 0  0.0 

Euonymus americana 1 4 20  0.8 

 

0 0 0  0.0 

Fraxinus americana  4 25  1.0 

 

0 0 0  0.0 

Ilex opaca 2 8 45  1.9 

 

0 0 0  0.0 



 

 

     

 

 

Mid-level 

 

FREQ RF #Stems MStems 

 

Canopy 

 

FREQ RF #Stems MStems 

Liquidambar styraciflua 13 54 385 16.0 

 

9 38 260 10.8 

Nyssa sylvatica 10 42 210  8.8 

 

2 8 45  1.9 

Ostrya virginiana 2 8 65  2.7 

 

0 0 0  0.0 

Quercus alba 8 33 220  9.2 

 

6 25 195  8.1 

Quercus lyrata 3 13 65  2.7 

 

7 29 330 13.8 

Quercus nigra 1 4 15  0.6 

 

2 8 60  2.5 

Quercus texana 0 0 0  0.0 

 

2 8 70  2.9 

Quercus phellos 3 13 25  1.0 

 

6 25 315 13.1 

Quercus rubra 3 13 70  2.9 

 

8 33 265 11.0 

Quercus sp. 1 4 15  0.6 

 

3 13 45  1.9 

Rhododendron canescens 2 8 35  1.5 

 

0 0 0  0.0 



58 

 

 

 

 

Mid-level 

 

FREQ RF #Stems MStems 

 

Canopy 

 

FREQ RF #Stems MStems 

Tilia americana 2 8 17  0.7 

 

0 0 0  0.0 

Ulmus serotina 2 8 45  2.7 

 

1 4 30 13.8 

Total Stems   2,087  

 

  1,710