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Invasion of Microstegium Vimineum (Poaceae), an Exotic, Annual, Shade-Tolerant, C4 Grass, Into a North Carolina Floodplain Author(S): Lawrence S

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Invasion of Microstegium Vimineum (Poaceae), an Exotic, Annual, Shade-Tolerant, C4 Grass, Into a North Carolina Floodplain Author(S): Lawrence S The University of Notre Dame Invasion of Microstegium vimineum (Poaceae), An Exotic, Annual, Shade-Tolerant, C4 Grass, into a North Carolina Floodplain Author(s): Lawrence S. Barden Source: American Midland Naturalist, Vol. 118, No. 1 (Jul., 1987), pp. 40-45 Published by: The University of Notre Dame Stable URL: http://www.jstor.org/stable/2425626 . Accessed: 29/09/2014 12:34 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Notre Dame is collaborating with JSTOR to digitize, preserve and extend access to American Midland Naturalist. http://www.jstor.org This content downloaded from 158.135.136.72 on Mon, 29 Sep 2014 12:34:07 PM All use subject to JSTOR Terms and Conditions Invasion of Microstegium vimineum (Poaceae), An Exotic, Annual, Shade-Tolerant, C4 Grass, into a North Carolina Floodplain LAWRENCE S. BARDEN DepartmentofBiology, University ofNorth Carolina at Charlotte,Charlotte 28223 ABSTRACT: Microstegiumvimineum, an Asian annual C4 grass that is veryshade- tolerant,has invaded floodplains,streambanks and adjacent mesic slopes in the NorthCarolina Piedmontduring the past 30 years. A 3-yearstudy of its invasive characteristicsrevealed that M. vimineumis slow to invadeundisturbed vegetation, but rapidlyfills disturbed, mesic, shaded areas, such as streamsideswhere floods scour existingvegetation or sewer-linerights-of-way which are mownonce a year.Its seeds remainviable forat least 3 yearsin the soil seedbankand rapidlygerminate to pro- duce a new cohortif a disturbanceremoves an existingcohort. On fertilefloodplain sites,soil fertilizationin March had no effecton seed productionin October.When M. vimineumseeds weresown into existing vegetation, seed productionwas negatively correlatedwith soil potassium,calcium, silt and pH, probablybecause the morefer- tile sitesalso supporteda denserground vegetation layer. These qualities,in addition to its cleistogamousor apomicticreproduction, help explain how M. vimineumhas spreadthroughout the eastern U.S. sinceits introduction approximately 70 yearsago. INTRODUCTION Microstegiumvimineum (Trin.) A. Camus is an annual grass species fromAsia which recentlyinvaded floodplains,streamsides and adjacent mesic slopes in the North Caro- lina Piedmont. It appears to be advancing into the existingground cover of floodplains, which consists mainly of Japanese honeysuckleLonicera japonica Thunberg. These two species often occur in dense monospecificstands which are separated by a strikingly narrow zone of mixing. This patternof an introducedannual grass invading established stands ofJapanese honeysuckle,a perennial woody vine which is known for its aggres- siveness on fertilesites of the Carolinas (Bruner and Shearin, 1964), led me to ask the followingquestions: (1) Is M. vimineuminvading undisturbed stands of L. japonica? (2) Does M. vimineumcreate a seedbank? (3) Can M. vimineuminvade undisturbedstands of L. japonica if its seeds are sown in the stands in large numbers? (4) What environmental variables are related to success of M. vimineumsown in stands of L. japonica? (5) Does M. vimineumrespond to an increase in essential soil elements with greaterreproductive success? (6) Does M. vimineumalter the soil so as to inhibit growthof L. japonica? (7) What is the role of disturbancein the invasion of M. vimineum? Microstegiumvimineum is in the tribe Andropogonae, subfamilyPanicoideae. It was firstcollected in the United States in 1919 near Knoxville, Tennessee (Fairbrothersand Gray, 1972). By 1933 it had been collected in the mountains of westernNorth Carolina (Blomquist, 1948; Fairbrothersand Gray, 1972) and by 1964 in 35 counties in North Carolina, primarilyin the Piedmont (Radford et al., 1968). By 1972 the species had spread to at least 14 eastern states, from Florida to New Jersey and W to Ohio and Mississippi (Fairbrothersand Gray, 1972), and by 1978 to Arkansas (Smith, 1978). In Asia the species occurs in Japan, Korea, China, Ryukyus, Formosa, Malaysia, India and the Caucasus (Ohwi, 1984). Microstegiumvimineum is unusual in that it is a C4 plant which, unlike typical C4 plants, is adapted to low lightconditions (Brown, 1977, Winter etal., 1982). Under net- ting which transmitted18% full sunlight,dry matter production was not significantly reduced fromproduction in full sunlight; at 5% full sunlight,growth was 17% of that at full sunlight(Winter et al., 1982). Apparently,the C4 pathway itselfhas no inherent disadvantages, compared to the C3 pathway, under conditions of low light (Pearcy, 1983; Pearcy and Troughton, 1975). 40 This content downloaded from 158.135.136.72 on Mon, 29 Sep 2014 12:34:07 PM All use subject to JSTOR Terms and Conditions 1987 BARDEN: SHADE-TOLERANT GRASS 41 Both cleistogamyand chasmogamy have been reported forMicrostegium vimineum in Japan, with all flowerson axillary racemes cleistogamous (Tanaka, 1975). Near Char- lotte, N.C., anthersare rarelyseen extendingfrom flowers and then only in ca. 10% of the racemes of plants grown where moisture and light are abundant. I have never ob- served anthers extendingfrom flowers of plants grown in heavy shade, nor have I ob- served stigmas extendingfrom flowers on any plants (Barden, pers. observ.). The study area was a floodplainwith a forestof Acernegundo L., Fraxinuspennsylvanica Marshall and Platanusoccidentalis L. located on the Ecological Reserve on the campus of the Universityof North Carolina at Charlotte. Ground vegetationwas primarilyLoni- cerajaponica and Microstegiumvimineum. In summer, light beneath the canopy ranged from3-12% fullsunlight; in April beforeleaf emergence in the canopy, lighton the for- est floorwas ca. 40% full sunlight.At the study area, M. vimineumbegins germinating in March, reaches 5-10 cm in height by the time of canopy leaf emergence in April, flowersin September and sheds its seeds in October. The alluvial soil of the study area was primarilysandy clay loam or sandy loam, with a relativelyhigh cation exchange capacity of 14.8 + 0.4 meq/100 g ( ? SE), base sat- uration of 82 ? 1% and pH of 5.2 + 0.06 on 40 experimental plots. Soil moisture (% dry weight)was 38 + 2.0% 1 week afterheavy rain and 23 + 1.5 % after4 rainlessweeks on 20 plots in the floodplain. METHODS Experiment1.-To check for natural invasion by Microstegiumvimineum into undis- turbed stands of Lonicerajaponica, 10 line intercepts, each 5 m long, were placed throughoutthe study area inJune 1982 across sharp interfacesbetween adjacent stands of M. vimineumand L. japonica. In July, 1983, 10 more interceptswere similarlyestab- lished. The vegetationalong the interceptswas not disturbed and cover of each species was measured to the nearest 10 cm when the lines were establishedand at the end of the experimentin 1985. Measurements in 1985 were made within 10 calendar days of ini- tial measurementsto minimize differencesin phenological developmentbetween years. Change in cover ofM. vimineumon the 20 lines was statisticallyanalyzed by paired t-test in which final cover was paired with initial cover. Experiment2. -A seedbank experimentwas begun in 1983 to determine how long Microstegiumvimineum seeds survived in the soil. An isolated 2-m2 stand of M. vimineum on an upland site 100 m fromthe floodplainwas chosen for studybecause floods could not transportnew seeds into it and because of the absence of Lonicerajaponica or other potential competitorson the site. A similar stand 25 m from the firstwas chosen in 1984 for a second seedbank test. The two stands of M. vimineum,which were along a footpathused daily during another study in 1977-1981, probably originatedfrom seeds accidentallytransported from the floodplain on clothing. Each year before seeds were produced, all plants in the two populations were counted and removed fromthe sites. Care was taken not to transportany new seeds into the seedbank sites. Experiment3. -Twenty lines, each 5 m long, were established in undisturbedLonicera japonica stands which contained no Microstegiumvimineum before the experiment. The firstand last 1-m section of each line was sown with 3 g of M. vimineumseeds (approxi- mately 2400 seeds m'1) in October 1982. The center section of each 5-m line was left unsown, as a control. The experimentwas designed to determinewhether M. vimineum could establish self-perpetuatingpopulations in undisturbed stands of L. japonica. Mi- crostegiumvimineum plants were counted during summers of 1983 and 1984, and seed heads were counted in October 1983 and 1984. The length of each seed spike was also measured in 1984 so that the number of seeds produced could be estimatedfrom an al- lometricregression equation. Experiment4.-Several soil, biotic and otherenvironmental factors were measured at each of the 40 lines where Microstegiumvimineum seeds were sown to determine which factorsmight be correlated with the relative success of each population in producing This content downloaded from 158.135.136.72 on Mon, 29 Sep 2014 12:34:07 PM All use subject to JSTOR Terms and Conditions 42 THE AMERICAN MIDLAND NATURALIST 118(1) seeds. The factorsmeasured were LAI of all species, cumulative photosyntheticallyac- tive
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