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PHYTOGEOGRAPHY OF THE THICKET, EAST TEXAS BIG MacRoberts and MacRoberts Michael Barbara H. R. Bog Research 740 Columbia^ Shreveport, Louisiana 71104, U.S.A. and Museum l-lerbarium, of Life Sciences Louisiana State University in Slireveport, Slireveport, Louisiana, 71 U.S.A. 15, 1 ABSTRACT We We examined the floristic relationship between the Big Thicket of east Texas and the remainder of North America. found that floristi- cally the Big Thicket overwhelmingly eastern and notably southeastern. There very western influence on is is little it. RESUMEN Hemos Hemos examinado las relaciones floristicas entre el Big Thicket del este de Texas y el resto de Norte America. encontrado que florlsticamente el Big Thicket es sorprendentemente oriental y notablemente del sureste. Hay muy poca influencia occidental en el. INTRODUCTION made Extraordinary claims have been regarding the ecology of the Big Thicket of east Texas. has been It described as exceptionally species rich, unusually diverse in plant communities, and a biological "crossroads" & between eastern and western floras (see Cozine 2004; MacRoberts MacRoberts 2004; Diggs et al. 2006 On for reviews and literature). the other hand, recent examinations of these claims have led to the conclu- sion that the Big Thicket is not a crossroads but is southeastern with levels of habitat diversity and species & richness comparable to other areas of the southeast (MacRoberts MacRoberts 2004; Diggs et 2006; al. MacRoberts 2007). As Diggs (2006:166) conclude "these conflicting views need examination." et al. et al. However, such an examination has been hampered by a lack of adequate floristic data. Until recently, the Big now Thicket flora has been understudied and undocumented (MacRoberts et 2002b), a problem largely al. overcome by inventory of the major part of the Big Thicket National Preserve (MacRoberts a floristic et al. Brown 2002b; et al. 2005, 2006a, 2006b, in prep.), by a reexamination of earlier plant collections from the Big Thicket National Preserve (L.E. Brown pers. comm.; Warren Pruess pers. comm.), and by the publica- and tion oi Atlas of the Vascular Plants of Texas (Turner et al. 2003) Illustrated Flora of East Texas (Diggs et al. much now 2006). Additionally, better distribution information on the flora of North America available is & USDA Meacham North America 1993-2006). Kartesz 1999; 2007; Flora of (e.g., Figure shows the location of the Big Thicket. While the exact geographic boundaries of the Big 1 Thicket are ill-defined, it is generally considered to consist at least of Newton, Jasper, Hardin, Tyler, Polk, and northern Liberty county, or an area of approximately 13,500 square kilometers (see Diggs et [2006] al. and Within for a discussion of the Big Thicket location a description of its ecology). this area is located the Big Thicket National Preserve, with 12 units ranging from 222 ha to 10,100 ha and totaling about 34,000 ha. Since the units were originally selected to represent the Big Thicket community structure and flora (Peacock 1994; Cozine 2004; Diggs et al. 2006), a floristic inventory of the units should constitute a fair sample of the area. As part of a continuing study of the flora of the West Gulf Coastal Plain, in this paper we examine the relationship between the Big Thicket and the remainder of North America north of Mexico. floristic METHODS Brown Using the vouchered of MacRoberts (2002b), (2005, 2006a, 2006b, in 1. floristic lists et al. et al. Warren we prep.), Pruess (pers. comm.), for the Big Thicket National Preserve, developed a master of the list - J. Bot. Res. Inst. Texas 1(2): 1149 1155.2007 1150 Journal of the Botanical Research Institute of Texas 1(2) Location of Big Thicket (hatched area) and Caddo Parish (stippled area). Fig. 1 . vascular plant species occurring in the Big Thicket region of east Texas. This consists of 1076 species. list Using Kartesz and Meacham (1999) and Flora North America (1993-2006), we plotted the North American of by and distribution of these species province, region. state, 2. Because Texas is a large state and because there is a major floristic ecotone across its center where & east and west meet (MacRoberts MacRoberts 2003; Diggs et al. 2006), we divided the state into 24 areas km approximately 180 on a side and using Turner et al. (2003) and Diggs et al. (2006), we plotted the dis- tribution of the species where possible in our master (see method above) across Texas to determine the list 1 relationship between the Big Thicket and other areas of Texas. This consists of 1035 species. floristic list Using Matos and Rudolph Brown Turner and Diggs 3. (1985), et al. (2002), et al. (2003) et al. (2006), we located species from the Big Thicket area (Newton, Jasper, Tyler, Hardin, Polk, and northern Liberty all We county) not in our master (see method above). combined this with our previous making a list 1 list list Meacham we sample of 1462 species. Using Kartesz and (1999) and the Flora North America (1993-2006), of We determined their North American distribution. then classified all species into one of five categories: 1) western: species that occur exclusively or predominantly in the west (the floristic divide between east and west is about 97 or 98 west longitude [MacRoberts et al. 2003]) (e.g., Loejlingia squarrosa Nutt.), 2) central: species that are predominantly in the central plains area (e.g., Tradescantia occidentalis (Britt.) Smyth), 3) MacRoberts and MacRoberts, Phytogeography of the Big Thicket, East Texas 1151 Fig. 2. Distribution of Big Thicket flora as percentage of total in each state, region, or province. universal: species that occur in virtually states and provinces in North America Athyriumjelix-femina all (e.g., (L.) Roth), 4) eastern-western: species that are not universals but which occur equally in both the east and in the west (e.g., Evolvulus sericeus Sw., Phyla nodiflora (L.) Greene), 5) eastern: species that are either exclusively found in the east or are predominantly in the east Xyris spp.)(this category includes West Gulf Coastal (e.g., Our Plain endemics [MacRoberts 2002a]). purpose was determine the influence of different regions et al. to on the flora of the Big Thicket. Brown Using MacRoberts 2006a, 2006b, Turner 4. et al. (2002b), et al. (2005, in prep.). et al. (2003) and Diggs et al. (2006), we located all species from the Big Thicket area (as in method 3) that also occurred Gameron in south Texas (Kenedy, Brooks, Jim Hogg, Zapata, Starr, Hidalgo, Willacy, counties) and deter- mined North American Our purpose was determine their distribution. to the distributional characteristics common between and what commonality of species in the Big Thicket outlying regions to see their was. The two have 303 common. areas species in we 3. In order to develop a comparative sample, using MacRoberts and MacRoberts (2006) applied km methods 1 and 3 to the flora of Gaddo Parish, Louisiana. Gaddo Parish is located about 200 north of the Big Thicket in the northwestern corner of Louisiana, in the center of the West Gulf Goastal Plain (Fig. We used a random sample of 417 species of the Gaddo Parish flora as in method and 1168 species as 1). 1, method in 3. Exotic species are excluded from of our data all sets. RESULTS Figure 2 shows the distribution of Big Thicket flora as percent of total species found in each state, province, shows shows or region. Figure 3 the distribution of Big Thicket flora as percentage across Texas. Table 1 1152 Journal of the Botanical Research Institute of Texas 1(2) Fig. 3. Distribution of Big Thicket flora as percentage across Texas: West Gulf Coastal Plain outlined in dashes. Big Thicket outlined in dots. how North America and Caddo shows the regions of relate to the Big Thicket Parish. Table 2 the relationship common between the species that occur in between the Big Thicket and south Texas. Figure 4 shows the Caddo distribution of Parish flora as percent of total species found in each state, province, or region. DISCUSSION 98% The method show results of that the Big Thicket phytogeographically eastern; notably southeastern: 1 is 88% 48% New of the Big Thicket flora occurs in Louisiana, 91% in Mississippi, in Florida and Georgia, in New common York, but only 19% in Mexico. In fact, the Big Thicket has more of its flora in with Wisconsin, New Michigan, Minnesota, Ontario, and England than has with any western state or province. Method it 2, not surprisingly, shows that within Texas the Big Thicket is floristically most similar to east and central Texas, notably to the West Gulf Coastal Plain region of which it is a part. Method 3 shows that eastern spe- cies dominate the Big Thicket, with 81% of the total, followed by eastern-western species, with 10%, and 96% universal species, with 5%. Western and central species account for slightly less than 4%; that over is, Method shows of species are eastern associates (eastern, universal, eastern-western). 4 that, of the species MacRoberts and MacRoberts, Phytogeography of the Big Thicket, East Texas 1153 and Caddo Table Regional affiliation of Big Thicket Parisli flora 1 . Caddo Big Thicket Parisli Number Number Region of Percent of of Percent of Species Species Species Species Western 19 17 1.3 1.4 Central 36 2.4 51 4.4 Universal 77 5.3 85 7.3 Eastern-Western 143 9.8 181 15.5 Eastern 1187 81.2 834 71.4 1462 1168 Total 100.0 100.0 common Table 2. Regional affiliations of species in to floras of Big Thicket and south Texas. Number Region of species Percent of Species Western 9 3.0 Central 12 4.0 Universal 33 10.9 Eastern-Western 102 33.7 Eastern 147 48.4 Total 303 100.0 common in between the Big Thicket and south Texas, eastern species dominate, with 48%, followed by 93% eastern-western species, with 34%, and universal species, with 11%; in other words, of the species are 7% eastern associates (eastern, universal, eastern-western). Western and central species account for only of common the total. This last analysis also makes clear that the vast majority of species held in between such areas as south Texas and the Big Thicket are eastern, not western, indicating that the Big Thicket not an is ecotone or "crossroads" between east and west but located fully in the eastern part of the North Ameri- is can region (Austroriparan subprovince) (McLaughlin 2007). Almost without exception western and floral central species that occur in the Big Thicket are very rare there and are usually associated with specialized uncommon and such and barrens Smith) Smith, habitat, as prairies Croptilon rigidijolium (E.B. E.B. (e.g., Hedeoma drummondii Benth.). The Caddo Parish analysis (method 5) reinforces this conclusion. While Caddo Parish has more in com- mon with states and provinces north of Arkansas, Oklahoma) than does the Big Thicket (MacRoberts it (e.g., & New New MacRoberts 2006, unpublished has approximately the same relationship with Mexico, data), it York, Mississippi and Florida as does the Big Thicket. Additionally, while Caddo Parish partakes of the same percentage of western species as does the Big Thicket, has a higher percentage of central, universal, and it West east-west species than does the Big Thicket. Clearly, therefore, the Big Thicket not exceptional in the is Gulf Coastal Plain in its affinities to other regions except perhaps the east and/or southeast. What many can be truthfully said about the Big Thicket that eastern plants reach their western and is Some southern limit in that region. of these species are: Cypripedium kentuckiensis C.F. Reed, Cy villa racemijlora Eriocaulon compressum Lam., Fagus grandijlora Ehrh., Hypericum galioides Lam., Lechea minor Leucothoe L., L., racemosa (L.) Gray, Rhexia petiolata Walt., Rhododendron canescens (Michx.) Sweet, Stewartia malacodendron Many and Chapman. L., Triantha racemosa (Walt.) Small, Trillium recurvatum Beck, Xyris stricta other eastern much species get farther west than the Big Thicket; examples include: Drosera capillaris Poir., Eriocaulon decangulare L., Liatris pycnostachya Michx., Rhexia mariana L., Sarracenia alata Wood, and Xyris torta Sm. Eew truly western or central species reach the Big Thicket; western and central habitat simply not there is very or limited. is 1154 Journal of the Botanical Research Institute of Texas 1(2) 4. Distribution of Caddo Parish flora as percentage of total in each state, region, or province. Fig. ACKNOWLEDGMENTS Warren Pruess provided an updated (January 20, 2007) of the plants collected from the Big Thicket list was by Agreement National Preserve, a project that supported, in part, a National Park Service Cooperative Two (No. 14001004) Paul Harcombe, Department of Ecology and Evolutionary Biology, Rice University. to anonymous made reviewers useful suggestions. REEERENCES Windham and The Brown, Hillhouse, B.R. MacRoberts, M.H. MacRoberts. 2002. vascular flora of Prairie, Polk L.E., K., County, east Texas. Texas 54:227-240. Sci. J. Brown, I.E., B.R. MacRoberts, M.H. MacRoberts, P.A. Harcombe, W.W. Pruess, LS. Elsik, and D. Johnson. 2005. Annotated checklist of the vascular flora oftheTurkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin 807-1 counties, Texas. Sida 21:1 827. W.W. and Annotated Brown, MacRoberts, M.H. MacRoberts, RA. Harcombe, 2006a. I.E., B.R. Pruess, I.S. Elsik, S.D. Jones. checklist of the vascular flora of the Big Sandy Creek Unit of the Big Thicket National Preserve, Polk County, 22:705-723. Texas. Sida W.W. and Annotated Brown, MacRoberts, M.H. MacRoberts, RA. Harcombe, 2006b. I.E., B.R. Pruess, I.S. Elsik, S.D. Jones. checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County, 175-1 Texas. Sida 22:1 189. W.W. and Annotated Brown, L.E., B.R. MacRoberts, M.H. MacRoberts, RA. Harcombe, Pruess, Elsik, S.B.Walker, in prep. I.S. checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. MacRoberts and MacRoberts, Phytogeography of the Big Thicket, East Texas 1155 CoziNE, 2004. Saving the Big Thicket: From exploration to preservation, 1685-2003. Univ. North Texas Press, J. Denton. and DiGGs, G.M., B.L. Lipscomb, M.D. Reed, RJ. O'Kennon. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26: 1-1594. Flora of North America Editorial Committee (eds.). 993-2006. Flora of North America North of Mexico. Vols. 2, 1 1 , 3, New 4,1 20, 21 22, 23, 25, 26. Oxford Univ. Press, York and Oxford. 9, , Kartesz, J.A. and CA. Meacham. 999. Synthesis of North American flora. Version North Carolina Botanica .0. 1 1 Garden. Chapel Hill. MacRoberts, M.H., B. R. MacRoberts, B. Sorrie, and R. Evans. 2002a. Endemism in the West Gulf Coastal Plain: impor- tance of xeric habitats. Sida 20:767-779. MacRoberts, B.R., M.H. MacRoberts, and I.E. Brown. 2002b. Annotated checklist of the vascular flora of the Hickory Creek Unit of the Big Thicket National Preserve,Tyler County, Texas. Sida 20:781-795. and The MacRoberts, M.H. B.R. MacRoberts. 2003. east-west transition of flora inTexas: a biogeographical analysis. 20:1693-1700. Sida and The MacRoberts, M.H. B.R. MacRoberts. 2004. Big Thicket: typical or atypical. East Texas Hist. Assoc. 42:42-51. MacRoberts, B.R. and M.H. MacRoberts. 2006. An updated, annotated vascular flora of Caddo Parish, Louisiana, on phytogeography and 191-1219. with notes regional ecology. Sida 22:1 and MacRoberts, M.H., B.R. MacRoberts, R.G. Kaunsky. 2007. Vascular plant species/area relationships (species rich- theWest ness) Gulf Coastal Plain:afirst approximation. Bot. Res. Texas 1:577-583. in Inst. J. Matos, and D.C. Rudolph. 1985. The vegetation of the Roy Larsen Sandylands Sanctuary the Big Thicket J.A. E. in ofTexas. Castanea 50:228-249. McLaughlin, S.R 2007. Tundra to tropics: the floristic plant geography of North America. Sida, Bot. Misc. 30. A&M Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas Press, College Station. and Turner, H. Nicols, G. Denny, 0. Doron. 2003. Atlas of the vascular plants ofTexas. Sida Bot. Misc. 24: B.L., 1-888. Usda 2007. The PLANTS Database (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA nrcs. 70874-4490.

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