AND COMPOSITION POTENTIAL COMPETITORS FLORISTIC IN ^ MEUSSIFOLIA (LAURACE COLONIES LINDERA AE) IN MISSISSIPPI WITH REFERENCE TO HYDROLOGIC REGIME Hawkins 2 Daniel Skojac 3 Nathan M. and Theodor Leininger 1 A. Schiff, D. Tracy - Jr. S. , , USDA Forest Service Center Bottomland Hardwoods Research for U.SA 3 Stoneville, Mississippi 776, 1 ABSTRACT fill RESUMEN INTRODUCTION Species member United This ^melissifolia (Walter) Blume a dioecious shrub endemic to the southeastern States. is *l»aceae, commonly was South Carolina and described by Walter (1788) called pondberry, collected in ® Uurus few herbarium specimens wtissaefolia. By the mid-20th century, Steyermark (1949) found that “ made dbeen having been collected subsequent with only three collections description, to Walter’s (1788) ^ 18 United throughout the first half of the twentieth Herbarium studies by Steyermark (1949) century. ^confirmed Missouri, Louisiana, Georgia, Alabama, Florida, the historical presence of in L. melissifolia Tennessee, T^Grolina, made in and South Carolina. Although Gattinger (1901) reference to L. melissifolia er® rk(1949) pubescent Lindera Benzoin, since la misidentified ^ believed this report probably referred “...to * Wtnentic University of Tennessee... herbarium of the material of has been found in the L. melissaefolia 7^’ with only one historic report (1924) excluded from “Shrubs of Indiana” citing L. melissifolia joucher concluded that Steyermark (1949) specimen. Based on the number of preserved L. melissifolia, ma Cies be ^ one United y of the rarest shrubs in the States. J^atly, Arkansas, Georgia, Mississippi, Alabama, extant present in populations of are L. melissifolia and In the Louisiana Florida. North be extirpated in Carolina, and South believed to Zj*"’ Carolina. It is 2** margin of seasonally grows along the Coastal ^ Plain (South Carolina and Georgia), L. melissifolia & (Alenc Brongn. de and Taxodium ascendens J*" pressional Walter wetlands dominated by Nyssa biflora depres- grows along the edge of a forested 2005). The Alabama known only population to occur in , 382 Journal of the Botanical Research Institute of Texas 4(1) Lower (LMAV), 2005), In the Mississippi Alluvial Valley L. melissifolia populations in Mississippi, Arkansas, and Missouri grow in periodically flooded, bottomland hardwood forests underlain by hydric soils. Forest LMAV canopy composition among composed the sites in the is similar, primarily of trees designated as facultative wetland or obligate wetland species (Hawkins et 2009a). Liquidambar al. styraciflua, Acer rubrum, Q. lyrata Q. nuttallii, and phellos are important canopy components; however, the relative importance Q. of these and other canopy species varies between sites (Hawkins et 2009a). al. Lindera melissifolia populations consist of spatially segregated, unisexual colonies (Hawkins etal. 2009b). Colony sizes range from approximately 20 stems to >1,000 stems (Morris 1987; Devall et 2001; Hawkins al. LMAV, pers. obs.). In the populations are strongly male-biased, with male female colony to ratios ranging from (Wright 1994) (Hawkins 7:1 to 19:1 et 2009b). al. In both male (pollen-bearing) and female (seed/fruit-bearing) L. melissifolia, anthesis occurs in bte spring, often during flooded conditions, and precedes leafing out. Small, yellow flowers are produced on axillary inflorescences (Fig. 1A), followed by production of green drupes on female plants. Approximately 90 days following anthesis, drupes contain a single, fully developed seed (Connor et 2007) and upon al. maturation drupes are bright red (Fig. IB, 1C). Drupes are dispersed from to early winter (Smith etal. fall and may 2004) fruit pedicels remain on the plant until the following spring. Female L. melissifolia appear to invest heavily in sexual reproduction (Connor et 2007); however, seedlings are rarely observed in al. naturally occurring populations (Wright 1990). Vegetative propagation of ramets from rhizomes appears to be the predominant form of reproduction (Wright 1990, 1994). During one the first to three years following seedling and/or ramet emergence, L. melissifolia plants are morphologically very similar to L. benzoin Blume. In during Steyermark’s (1949) investigation (L.) fact, of the species, he noted numerous misidentified herbarium specimens of resulting from this L. melissifolia close similarity. Although leaf pubescence and leaf size sometimes differ between the two species, these may characteristics be dependent on season and locality. On the other hand, the angle of the lateral veins may in the leaf blade be used to distinguish the congeners. In the lowest two pairs of lateral L. melissifolia, veins of the leaf blade diverge at a 45°-50° angle from the midrib, and successive veins diverge distal to this, at approximately 35° to the (Steyermark, 1949). In other words, the lower two veins are not parallel lateral successive upper same veins. In contrast, all lateral veins of L. benzoin leaves diverge from the midrib at the (194#*r angle (35°-45°); therefore, are parallel (Steyermark Both and Steyermark 1949). Nuttall (1818) scribed the fruits of L. melissifolia as “larger” than those of L. benzoin. The senior author has found the seeds L of the congeners be While seeds of to consistently reliable for definitive identification of fruiting plants. biow» melissifolia are spherical and brown brown and dark light to yellowish those of L benzoin are oval (Fig. ID). and In 1986, was Fish L. melissifolia listed under the Federal Endangered Species Act of 1973 (U.S. (Hawknwjw^ Wildlife Service 1986). Recent research has provided some ecophysiology insight into the & wi*T 2009b; Kirkman Aleric 2005) of and associated this species, types and structure forest forest LMAV speck* melissifolia in the have been suggested reported (Hawkins Wright (1990) 2009a). et al. peri** Brunnichia, Rubus, and Smilax main and that as competitors Arkansas, of northeast L. melissifolia in flooding served very minimize to competitive Beyond report, interactions. Wright’s (1990) initial P^ known about is the to microhabitat or biotic interactions in colonies. In an effort L. melissifolia concise information disjo** regarding native, the microhabitat and three general ecology of L. melissifolia, populations asse® in Mississippi were monitored for three years. The objectives of our study were to 1) vascular plant P*#** checklist of groundcover identify species growing within colonies, 2) L. melissifolia competitors of L. and melissifolia, monitor 3) hydrologic regime colonies. for L. melissifolia Lindera melissifolia colonies L, and seeds L. melissifolia, (D) ! 384 Journal of the Botanical Research Institute of Texas 4<1) METHODS Study Sites Two The North of the study are in Sharkey County, Mississippi. Delta National Forest (NDNF) sites site a is 25-ha tract of Delta National Forest that has been under management by the U. S. Forest Service since 1938 & Ramp (Devall 1992). The South Delta National Forest (SDNF) site is a 10-ha tract of forest approximately km NDNF. 9 southeast of The third study site (BC) is in Bolivar County, Mississippi, and is a privately owned 30-ha fragment surrounded by Hawkins 2009a map and forest agricultural fields (see et al. for detailed Dowling descriptions). Soil association for the three sites is (very fine, smectitic, nonacid, thermic Venic Chromic Endoaquepts)-AUigator thermic Dystraquerts)-Sharkey (very-fine, smectitic, (very-fine, smectitic, thermic Chromic Epiaquerts), an association characterized by poorly drained, fine-textured clayey surface & and formed from Mean soils subsoils Mississippi River alluvium (Rogers 1958, Scott Carter 1962). annual temperature for Sharkey and Bolivar counties approximately 18.0°C, and total annual precipitation ranges is mm mm & from 1228 1319 to (Rogers 1958, Scott Carter 1962). Data and Collection Analysis autumn BC * NDNF In 2003, rectangular plots were established for selected L. melissifolia colonies at (Ncoiony 6), = and SDNF = SDNF was numerous many ( Ncoiony 10), (Ncoiony 1). In 2000, habitat for L. melissifolia colonies, with >200 stems (GSRC 2000). However, during reconnaissance of this area we found only one remaining lm colony and were able to establish only one study plot. The perimeter of each plot was positioned beyond m A cm the outermost L. melissifolia stems of a colony. 1.2 wooden stake x 5 cm) was established at each (5 comers of the four of the rectangular plot. Colony sizes were variable, and thus plot areas varied with colony lm lm size. Within each plot, x quadrats were marked with pin along the diagonals of the rectangular flags plot; therefore, the percent of plot area sampled was the same among plots. In June 2004, May 2005, and June 2007, L. melissifolia stems in each lm x lm quadrat within the {Jots Common were and were counted, groundcover species were and stems were counted. plants identified TSH When were identified in the field by and DAS. was problematic, plants in question field identification top-cropped and by Godfrey taken supplemented to the lab for identification using Radford et (1968), al. and Wooten (1979, 1981). From 10 November 2004 22 May When were not Hooded, to 2007, plots were visited bi-weekly. plots mewl two samples were cm soil collected from each plot and placed in individual 141 3 hermetically sealed, , an oven at containers. Soil samples were taken directly to the lab, weighed to the nearest 0.01 dried in g, 70°C for 48 hr, then weighed again. Percent moisture each sample was calculated by dividing the for soil W- by difference of and quotient initial (wet) final (dry) weights by the weight and multiplying the initial and During comer post flooded cm conditions, water depth was measured each plot to the nearest 0.1 at mean the SE) used (± to represent water depth for the plot. NDNF, colonies At were there substantial differences in water depth among some of the L. melissifolia mean analysis one-way therefore, water levels for each sampling date each colony were compared using a at of variance (ANOVA; SAS different (p 2001). For colonies where mean water was not significantly level 0.2421), data were pooled producing one mean ± standard error for each of three groups of colonies. AND RESULTS DISCUSSION defini- Hydrologic broad regime for forests with Lindera melissifolia populations often described within the is tion of “periodically flooded”. For populations occurred in in Mississippi, flooding generally BC The -spring. vever, flooding events and among flood duration were found vary sites. to d and annual flooding events of comparable depth and duration in 2005, 2006, 2 ^ regime t this site is artificially controlled by the lan refore, hydrologic i from isistent year to year. . m #0* Flood duration and time NDNF. occurred initial of flooding varied with year Flooding at c 2006, but not in 2007 monitored i (Fig. 2B). Although water depth differed among some of the ten NDNF, had no influence on time and duration of individual colony inundation this (Fig. 2B). Difference in among water depth colonies is the result of surface topography (e.g. sinks or sloughs). Lack of difference in time and duration of flooding between colonies results from rapid rising and dropping of water. In contrast NDNF SDNF to and BC, did not experience flooding in 2005, 2006, and 2007 (Fig. 3). When L. melissifolia colonies were not flooded, soil moisture content was comparable among the three 20% 30% sites, ranging from to throughout the year, with the exception of mid- to late-summer, when soil NDNF moisture contents dropped as low as 15% to 18% for BC and (Figs. 2A and 2B); and as low as 10% SDNF (Fig at 3). The combined annotated list of vascular plants (including L. melissifolia for the three study sites includes ) 70 species in 57 genera in 45 families (Appendix and of these, 9 species have the potential to become 1) weedy or invasive (SWSS 1998). The contribution of L. melissifolia to overall colony composition ranged from 20%-40% NDNF 5%-15% SDNF approximately at and BC, and at (Fig. Other species growing within the 4). colonies are typical of bottomland hardwood forests in this area of the LMAV, and tend to reflect hydrologic regime at each study site. Forty-nine percent of the 69 species identified as growing in association with L. FACW 67%-99% melissifolia (Appendix have a wetland indicator status of and a probability of occurrence 1) NRCS SDNF wetland (USDA, in a area 2008). However, the presence of Callicarpa americana at is atypical LMAV for bottomland forests in the and represented a county record reported in 2007 (Skojac et al. first SDNF Lack 2007). of inundation at (Fig. has possibly allowed establishment of C. americana, as well as 3) other species, such as Asplenium platyneuron and Phytolacca americana, that generally are not found in forests prone to flooding. Of 69 immediate the species growing in association with few appeared to pose an L. melissifolia, above-ground competitive threat and the ratio of stem density for these plants to L. melissifolia stem density remained relatively stable throughout our study The predominant growth habit in L. melissifolia (Fig. 4). colonies was vines (Fig 4). Wright (1990) considered Brunnichia ovata as a plant with potential to be an ag- gressive competitor of L. however, we observed early-summer emergence of B. ovata, as well as melissifolia; Toxicodendron and radicans radicans, after L. melissifolia plants had flowered and leafed out. Both B. ovata T. remained prostrate throughout the season and did not compromise capture by L. melissifolia leaves, light nor twine around or climb On bona-nox, glauca, S. L. melissifolia stems. the other hand, Smilax spp. (S. S. become rotundifolia, S. tamnoides) and Vitis spp. palmata, have potential to (V. aestivalis, V. V. rotundifolia ) stems strong competitors, by remaining upright throughout and some using melissifolia the year, in cases, L. above-ground for support. Many have direct of the associated species in do not appear to L. colonies in Mississippi melissifolia those competitive impact on and/or growth habit L. populations. However, with vining melissifolia species a response with potential become weedy The to or invasive (Appendix should continue to be monitored. 1), interactions of these species to natural or anthropogenic competitive disturbances has the potential to alter within these L. melissifolia populations. APPENDIX 1 (Monilophytes Plants included in this checklist are compiled in alphabetical order by family within two major groups names Angiosperms). Genera and common species are alphabetical within each nomenclature and family. Scientific (USDAN The Plants Database (USDA, NRCS Database 2008). Family circumscriptions monilophytes follow The Plants for aredeno^ and 2008), for angiosperms, APG 998) (Stevens become weedy (SWSS, 2008). Plants with potential to or invasive 1 with an asterisk (*) before the species name. Region (AL AR, Species wetland indicator status for the Southeast (67%'W*^ NC,SC,TN) = noted obl wetland is as: obligate (99% = probability ofoccurrence wetlands); facw facultative in ability of occurrence <* in wetlands, but occasionally found = (equal probability in non-wetlands); fac facultative m wetlands w or non-wetlands); = facu facultative upland (occurrence non-wetlands, occasional usually in and - - ni unable to determine wetland indicator status based solely on genus. Locality data are noted as: (1) BotlV M- - MS; 2 North County. ( ) Delta National Forest, Sharkey County, MS; and 3 = South Delta National Forest, Sharkey ( ) Journal of the Botanical Research 388 Institute of Texas4(1) Trees/Shrubs Vines Pondberrv Forbs/Herbs KS | BC -NDNF SDNF 1 1 per unit area (stems/m2 ). Melothria pendula L, Guadeloupe ci Cyperaceae Carex crus-corvi, Shuttlw. ex Kunze, r coralbead, fac, (2,3) Carex LH, Carolina louisianica Bailey, Louisiar *Cocculus carolinus DC., (L.) Carex tribulokles Wahlenb., blunt brc Moraceae swamp obl (2) privet, Forestiera acuminata (Michx.) Poir., 33) Marsh, green ash, facw, (1 Fraxinus pennsylvanica pumpkin ash, obl, • Fraxinus profunda (Bush) Bush, Gleditsia triacanthos L, honeylocust, (13 fac, Ludwigia glandulosa Walter, Fagaceae Quercus lyrata Walter, overcup oak, (U) obl, Orchidac Quercus nigra water n oak, L., fac, (3) tresses, October lady's ovalis Lindl, ?s Quercus pheltos L, willow oak, facw, (1,23) Quercus texana Buckley, Texas red oak, obl, (1 passionflower, fa yellow *Passifiora lutea L, Uquidambar Phytolaccaceae sweetgum, styraciflua L, fac, (1, pokeweed, f* * American Phytolacca americ i L„ Carya aquatica (Michx. Nutt, water f) hickor 389 Smilax glauca Walter, cat greenbrier, fac, i ovata (Walter) Shinners, redvine, facw, Smilax rotundifolia roundleaf greenbrk unnichia (1,2,3) L. 'Sr Polygonum virginianum L, jumpseed, fa Smilax tamnoides L, bristly greenbrier, fai (2) c, Styracaceae d leatherflower, facw, Styrax americanus Lam, American snowbell, facw, (2) (2) Ulmaceae supplejack, facw, Planera aquatica Gmel, water elm, obl, J.F. (2) i Ulmus americana American elm, L, facw, (1 ,2,3) Rubus Michx, Boehmeria cylindrica Sw, false nettle, facw, (23) trivialis (L.) buttonbush, Callicarpa americana L, American beautyberry, facu, obl, (3) (2) Violaceae Viola sp. L, violet, ni, (3) *Ampelopsis arborea (L) Koehne, peppervine, fac, (133) *Parthenocissus quinquefolia Planch, Virginia creeper. (L.) fac, (1,23) summer Michx. grape, Vitis aestivalis fac, (2) palmata Vahi, catbird grape, facw, Vitis (1,2) Michx, muscadine, rotundifolia fac, (23) Vitis ACKNOWLEDGMENTS Ihe authors thank Greg Comer, Stephanie Skojac, and Theran Stautz for their assistance in the field, U. S, Fish and Wildlife Service for permits, the U. S. Army Corps of Engineers for underwriting the cost of this research, and Drs. Charles Bryson, Emile Gardiner, and Edward W. Chester for review of an earlier draft °f this manuscript. REFERENCES Alewc endangered K-M. and LK. Kirkman. 2005. Growth and photosynthetic responses of the federally shrub, ' Lindera Amer. 92:682-689. melissifolia (Lauraceae), to varied light environments. Bot. J. ^ Cqnn and! K, G5. 2007. Schaefer, Donahoo, M. Gardiner! Hawkins, D. Wilson, N. Schiff, Hamel, Leininger. Devall, P. J. E. Development, endangered pondberry fatty acid composition, and storage of drupes and seeds from the (Undera melissifolia). Cons. 137:489-496. Biol. DtAM.CC. 1924. Shrubs of Indiana. Indiana Department of Conservation, Indianapolis. Pub. No. 44. Dewu MS. and and protection of old growth in the » P.F. Ramp. 1992. U.S. Forest Service research natural areas Nat. Areas. 2:75-85. J. 1 N endangered pondberry, Undera Schiff, and D. Boyette 2001 Ecology and reproductive biology of the - . Melissifolia (Walt.) Blume. Nat. Areas 21 :250-258. J. *®* A I1 - 190! The flora of Tennessee and a philosophy of botany, respectfully dedicated to the citizens of enn essee.„ Press of Gospel Advocate Publishing Co, Nashville, TN. R k and United Monocotyle- - J.W. Wooten. 979. Aquatic and wetland plants of the southeastern States. 1 5*** Unive rsity of Georgia Athens. Press, ^ UK. and J.W. Wooten. 198! Aquatic and wetland plants of the southeastern United States. Dicotyledons, diversity of Georgia Athens, GA. Press, South pondberry Prepared Research Corporation (GSRQ. 2000. survey report: reevaluation of in Mississippi. Final US. Army Corps of Engineers, Vicksburg Gulf South Research Corp, Baton Rouge, LA. ^ District, DS. Bottomland the and 2009a. forests in Skojac, B.R. Lockhart, T.D. Leininger, M.S. Devall, NJVI. Schiff. UJvver Mississippi endangered Undera Castanea 74:105-1 13. with the melissifolia. Alluvial Valley associated