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Evaluating the quality of a disturbed wetland in southwestern Indiana: a survey of native and exotic flora at Vectren Conservation Park.

ABSTRACT. Wetland stability promotes ecosystem services such as water purification and maintenance of biodiversity. These ecosystem services have been disrupted by anthropogenic degradation of natural habitats resulting in decreased biodiversity and the spread of introduced species. In Indiana, more than 87% of wetlands have been destroyed or degraded; those wetlands that remain are threatened by invasive species. To assess the need for restoration at Vectren Conservation Park in Southwest Indiana, a survey of the floral species present, as well as a study of the relative abundance of native and exotic species, was performed. The site includes more than 1100 acres of wetland habitat, including riparian forest, recently planted trees, and abandoned agricultural land. We collected 144 species from 109 genera, with 31 of the species being non-native to Indiana. When including all native and non-native species, the floristic quality index (FQI) of the site was 23.5 and the mean coefficient of conservatism ([C.sub.av]) was 2.0. The FQI and mean coefficient of conservatism ([C.sub.av]) were relatively low compared to other sites found in Indiana, indicating few natural remnants remain at the site. Although highly degraded, the site is capable of supporting high quality native wetland species, which would result in the improvement of ecosystem services and buffer against more extensive establishment of non-native species.

Keywords: coefficient of conservatism, floristic quality index, invasive species, restoration, species diversity, wetlands

INTRODUCTION

Ecosystem stability contributes to processes such as water purification, flood control, ground water recharge, and even maintenance of biodiversity. Anthropogenic degradation and destruction of natural habitats negatively impacts ecosystem stability (Pearson 1972; Van Auken 2000), which in turn threatens biodiversity and creates opportunities for colonization by invasive species (Burke & Grime 1996). In the United States, more than 50% of wetlands (Nichols 1988; McCorvie & Lant 1993) and up to 99% of prairies (Samson & Knopf 1994; Samson et al. 2004) have been destroyed or degraded, resulting in imperilment of native species.

Because so little high quality native habitat remains in Indiana, restoration of degraded habitats must be a priority. Efforts aimed at restoring degraded habits have become more widespread as awareness regarding the benefits of natural habitats has increased (Rood et al. 2003; Clark 2003). Restoration can reestablish ecosystem services in degraded habitats (Gratton & Denno 2006; Benayas et al. 2009), as well as enhance biodiversity. In turn, biodiversity can have a positive feedback on ecosystem function (Vernberg 1993; Lehman & Tilman 2000; Benayas et al. 2009). For example, diverse wetland plant communities purify water and provide flood control (Vernberg 1993; Benayas et al. 2009). Increasing diversity may also buffer against species invasions (Naeem et al. 2000; Kennedy et al. 2002, but see Foster et al. 2002; Eriksson et al. 2006).

After habitat destruction, biological invasions represent the second greatest threat to biodiversity (Vitousek et al. 1996; Knight & Reich 2005), so enhancing native diversity needs to be a priority when performing restoration ecology. Invasive species diminish biodiversity by outcompeting and excluding native species resulting in homogenization of habitats (Kaufman 1992). In some U.S. states, up to 47% of the flora is composed of exotic species. (Rejmanek & Randall 1994; Vitousek et al. 1997). In Indiana, invasive species pose a significant threat to the state's native flora (Weber & Gibson 2007), with approximately 39% of the 2800-plus species occurring in Indiana being categorized as non-native (Rothrock & Homoya 2005)

Special attention should be paid to biodiversity when restoring Indiana wetland habitat. First, wetlands are of particular interest when considering restoration because they support diverse habitats containing many rare species; in Indiana, wetlands lay claim to some of the highest levels of species diversity of any ecosystem type (Myers 1997). Second, more than 87% of Indiana's wetlands have been drained or destroyed (Dahl 1990; Miller & MacGowan 2004). Finally, wetland areas are especially susceptible to invasions; nearly 25% of the world's most invasive plants occur in wetlands (Zedler & Kercher 2004).

Vectren Conservation Park (VCP), a wetland next to the Wabash River in southern Indiana, may be a candidate for conservation or restoration. Comprised of abandoned agricultural fields and riparian forests, qualitative and quantitative surveys of native and exotic flora are necessary to evaluate the quality of the habitat in order to make conservation and restoration decisions for the site. One measure of plant diversity, species richness (i.e., the number of species in an area), provides a means to evaluate ecosystem quality. Examination of native versus exotic plants provides a second assessment tool. In this study, we performed a qualitative survey of the property to identify the plant species at the site, as well as conducting a quantitative study to determine the relative abundance of native and exotic species at the site. In addition, to provide metrics of floristic and ecological quality comparable to other Indiana sites, a floristic quality index (FQI) and a coefficient of conservatism (C) were generated for the site.

METHODS

Study Site.--VCP is an l 118 acre property located in southwestern Indiana (38[degrees]17' N, 87[degrees]52'W) (Fig. 1). Found 6.75 miles north-northeast of the town of Griffin in Posey County, the property is approximately 380 feet above sea level. In 2007, Vectren Corporation provided the University of Evansville (UE) with a long-term lease to the property in order to provide a research site for undergraduate students and UE faculty. Surrounded on three sides by the Wabash River, VCP regularly floods, occasionally being entirely inundated with water. The soil composition of the site varies. (McWilliams 1989). The soils at the northern end of the property are characterized as frequently flooded silt loams, while the southern half of the property includes a wide range of frequently and occasionally flooded silt loams, silty clay loams, and fine sandy loams. The site consists of 157 acres of riparian forest, 454 acres of recently restored forest (see below for details), and 508 acres of meadow (Woodburn 2001). In addition, an operating agricultural field of 81 acres exists in the middle of the meadow. The remains of a levee erected by farmers follow the path of the river, and most of the mature riparian forest is bounded by the farmer's levee. The meadow resides in the interior of the site.

Based on U.S. census data, most of VCP was farmed from the early 1800's until 2001 when the Vectren Corporation purchased the property. In 2002, Vectren Corporation planted 454 acres with trees and shrubs, including five species of oak and three species of dogwood, as well as sycamore, black walnut, sweetgum, spicebush, and button bush (see Appendix 1 for a complete list of species planted and the number of each species planted). The trees and bushes were purchased from Vallonia State Nursery in Vallonia, Indiana. Of the trees planted by Vectren Corp., sweetgum, black walnut, and sycamore have had the most success establishing at the site. In addition to the 136,100 trees and shrubs that were planted, selected areas including roadway easements and the area surrounding the 81-acre agricultural field, were planted with warm and cool season grasses. Except for the 81-acre agricultural plot, the land has remained mostly unmodified by human activity after Vectren planted the trees, shrubs and grasses.

Plant Survey.--In 2007, we initiated a survey of the flora of VCP. On a semi-weekly basis from May to October in 2007 and 2009, trips were made to VCP to document the flora present, both in the meadow and the forest. The recently replanted forests were avoided in the surveys. During each visit to the site, surveyors collected, pressed, and later identified any previously unidentified plant that was observed. The collected species were identified using a variety of identification keys and field guides (Deam 1940; Steyermark 1963; Gleason & Cronquist 1991; Holmgren 1998; Yatskievych 1999, 2006; Yatskievych 2000). The nomenclature from Gleason and Cronquist (1991) is reported for all species. The voucher specimens are being held at UE's herbarium.

Using plant information from the survey, each species was rated for level of wetland habitat preference, with categories including obligate and facultative wetland plants, as well as categories related to preference for upland habitat (U.S. Army Corps of Engineers 1996; see Appendix 2 for more details). The wetness scale proposed by Swink and Wilhelm (1994) was used to calculate a mean wetness value for VCP. In this scale, OBL = -5, FACW = -2.5, FAC = 0, FACU = 2.5, and UPL = 5; a site with a mean wetness below 0 is considered predominantly to have wetland plants.

In addition, each species was given a coefficient of conservatism (C-value) for Indiana (Rothrock 2004). C-values range from zero to 10, with lower values representing plants that are highly tolerant of disturbance and higher values representing plants that are usually restricted to high quality plant community remnants. Introduced plants are often not categorized for C-values, but they can be considered to have C-values of zero. A mean C-value (Cav) was calculated for the native species at the site. In addition, a Car for the combined values of native species and introduced species was calculated (after attributing a C-value of zero to the introduced species). To provide a floristic quality index (FQI), the two Car's were multiplied by the square root of the number of plant species.

Plot Samples.--In addition to the qualitative survey, a study examining the relative abundance of native and exotic plants occurring in meadow and forest environments was conducted during the summer of 2007. Three 20 X 20 m plots were randomly selected both in the meadow and in the forest habitat. The plots were constructed at least 100 meters from any farmlands, tree plantings, or access roads to minimize the effects of adjacent small-scale habitats. Within each plot, 12 subplots (0.5 X 1.0 m) were randomly sampled. If the subplot chosen included a large tree (whether standing or fallen), the subplot was moved to the next 0.5 X 1.0 m location directly to the left. In the forest, if a randomly selected subplot contained high concentrations of Toxicodendron radicans (poison ivy), the plot was moved to an area relatively free of this species. All forest plots were on the eastern edge of VCP. For all plots, ramet density was calculated by counting the number of stems of each plant species encountered in each subplot. Each species was identified as either native or non-native to the United States and to Indiana. Any species that could not be identified to species was identified to the lowest taxonomic level possible.

RESULTS

Plant Survey.--In total, 144 species from 109 genera within 54 families were collected from VCP (Appendix 2). Of the 144 species, 113 were native species and 31 were introduced species. Eight species were collected but identified only to genus because of a lack of morphological characteristics; these species were not classified as native or invasive. Seven additional species could not be identified to genus. Of those seven, one was a member of the Brassicaceae family, one was a member of the Poaceae family, and one was a member of the Apiaceae family. The final four species could not be attributed to a family because of a lack of identifying features.

Of the 113 native species at our site, twelve species (Asclepias incarnata, Senecio glabellus, Rorippa palustris, Rorippa sessiliflora, Cephalanthus occidentalis, Hibiscus laevis, Salix nigra, Phyla lanceolata, Forestiera acuminata, Amorpha fruticosa, Saururus cernuus, and Mimulus alatus) are almost always found in wetlands in our region. Overall, 42% of the species occur naturally at higher frequency in wetland habitat (FACW or OBL), and 36% are found equally as often in wetland habitats as in non-wetland habitats (FAC) (seed Appendix 2 for notation). The remaining 22% of native species are more commonly associated with upland habitats (FACU or UPL). The non-native species were plants less commonly associated with wetlands; 57% of the non-native species that had a wetland designation are more commonly found in upland habitat (FACU or UPL). Using Swink and Wilhelm's wetness scale (1994), the plants at VCP had a mean wetness value of -0.37, suggesting that the site is only weakly associated with wetland plants.

Few species had both high coefficients of conservatism (C-values of six or greater) and high fidelity to wetland habitats. The exceptions included Aristolochia tomentosa, Aster praealtus, Carex conjuncta, Celtis laevigata, and Forestiera acuminata (see Appendix 2 for details). Of the 107 native species with C-values, 49.5% had C-values of 0-2, and 43.9% had C-values of 3-5. The mean C-value ([C.sub.av]) for the native species was 2.5, and the [C.sub.av] for all species (native and non-native species combined) was 2.0. The FQI for native species was 26.6, while the FQI for all species was 23.5.

Plot Samples.--The composition of the three forest plots and the three meadow plots differed greatly. In general, woody species were more abundant in the forest plots, while the meadow plots were dominated by herbaceous species. In total, 17 families and 21 genera were represented in the forest plots, while 14 families and 26 genera of plants occurred in the meadow plots. Four families (Apiaceae, Cyperaceae, Poaceae, and Urticaceae) characterized the forest plots, accounting for over 91% of the species collected. The families characterizing the meadow plots differed from those of the forest; three families (Asteraceae, Fabaceae, and Poaceae) accounted for over 88% of the species in the meadow. Of the 23 species collected in the forest plots, only one was introduced (although four species were unidentified). Of the 29 species collected in the meadow plots, eight species were introduced (although four species were identified only to genus, and thus not classified as native or introduced). In the meadow plots, invasive species accounted for the majority of the stems counted (59.8%) (Table 1A). Together, Medicago lupulina (black medic), Melilotus officinalis (yellow sweet clover), Melilotus albus (white sweet clover), and Sorghum halepense (Johnson grass) accounted for almost all of the introduced stems counted in the meadow plots (98.4%).

In Meadow Plot I, native species accounted for almost all of the stems (98.1%). The plot was characterized by Aster praealtus (56.8% of the stems) and Elymus virginicus (25.6%). The majority of stems in Plot II belonged to introduced species, with Melilotus albus and Melilotus officinalis accounting for over 51% of the stems. The native species in Plot II primarily consisted of Aster praealtus (14.5% of the stems) and Solidago canadensis (15.7%). Plot III was largely characterized by introduced species, with Medicago lupulina accounting for over 70% of the stems. Oenothera biennis (10.2% of the stems) was the only native species commonly found in Plot III. Some species were found throughout all three plots, while others were clustered in one or two of the three plots. For example, all but one specimen of Medicago lupulina was found in Plot III, while most of the Melilotus stems were found in Plot II. Sorghum halepense was the only introduced species commonly found throughout the three meadow plots. The native species commonly found in all three plots included Aster praealtus, Solidago canadensis, and Elymus virginicus.

In contrast to the meadow plots, native species represented almost all of the species present in the forest plots. In fact, only two introduced specimens were found across the three plots--two specimens of Ipomea hederecea (ivy-leaved morning glory) in forest Plot I (Table 1B). Five species (Cryptotaenia canadensis (honewort), Carex grayi (globe sedge), Elymus virginicus (Virginia wildrye), Laportea canadensis (wood nettle), Pilea pumila (clear-weed), and Parthenocissus quinquefolia (Virginia creeper)) accounted for more than 95% of the stems counted in the forest plots. The most common species in forest plots were fairly consistent across plots. Forest Plot I was represented by Cryptotaenia canadensis (40.5%), Elymus virginicus (28.8%), and Pilea pumila (18.6%). Elymus virginicus (55.3%), Cryptotaenia canadensis (17.9%), Carex grayi (12.8%) accounted for most of the stems found in forest Plot II. In Plot III, Elymus virginicus (43.9%), Carex grayi (24.6%), and Pilea pumila (23.9%) were the most common species.

DISCUSSION

Plant Survey.--The quality of habitat ([C.sub.av] = 2.5, FQI = 26.6) at VCP is relatively low compared to other wetland sites in Indiana. For example, Bennett Wetland Complex (BWC) in Henry County has a [C.sub.av] for native species of 3.8 (Ruch et al. 2009) and an FQI of 54.6. Turkey Run State Park in Parke County, Indiana also contains two habitats of higher quality. Two seep areas in the park are categorized as high quality habitats ([C.sub.av1] = 5.4, [FQI.sub.1] = 29.8, [C.sub.av2] = 5.1 [FQI.sub.2] = 32.1) (Rothrock & Homoya 2005). The [C.sub.av] of VCP fails to fall within the suggested range of values for a natural habitat (>3.5), indicating that few natural remnants remain at this site. Although the site is degraded, it is still capable of sustaining species that require a wetland habitat. This is evident by the presence of Aristolochia tomentosa, Aster praealtus, Carex conjuncta, Forestiera acuminaa, and Celtis laevigata, which all have C-values equal to or greater than six and are commonly found in wetlands.

Plot Samples.--The proportion of non-native species in the meadow and forest plots differed greatly. The meadow has been used for agricultural purposes as recently as 2001, so the higher the abundance of invasive species was expected because disturbance promotes colonization by non-native species (Burke & Grime 1996). Although the invasive species accounted for over 50% of stems present in meadow plots, the presence of some non-native species (e.g., Medicago lupulina and Melilotus spp.) was localized, suggesting that concentrated efforts to control these non-natives may be successful.

In contrast, the widespread presence of Johnson grass, which was found in all plots, represents a pressing concern. Johnson grass produces large quantities of viable seed, while also spreading rhizomatously (Oyer et al. 1959). Furthermore, the formation of rhizomes as early as 50 days after seed planting causes the plants to become increasingly difficult to control because the entire rhizomatous system must be eradicated as opposed to simply destroying the aerial foliage. In addition, large quantities of herbicide are generally required to control Johnson grass (Frans et al. 1991), and some biotypes of Johnson grass have become resistant to certain herbicides (Smeda et al. 1997). For these reasons Johnson grass presents a major challenge in site restoration.

Restoration efforts have demonstrated that reestablishing biodiversity and ecosystem services can be effective. While VCP is a degraded site, restoring this environment could lead to improved ecosystem function, enhanced biodiversity, and reduced abundance of non-native plants (Vernberg 1993; Lehman & Tilman 2000; Benayas et al. 2009). Waterways, such as the Wabash River that borders VCP, expedite the spread of invasive species by acting as corridors for dispersal (Thebaud & Debussche 1991; Parendes & Jones 2000). So yearly floods currently wash in invasive seed banks and receding waters export invasive seed from the property's established invasive populations. In addition to restoring the ecosystem of VCP, establishment of a stable native wetland should result in exportation of native seed instead of non-native seed. For example, a few recently discovered species (i.e., Rudbeckia laciniata and Vernonia gigantea) may have come from an attempted prairie planting in 2008, where floods swept the seeds far from the planting.

APPENDIX 1:
Species planted by Vectren Corporation at Vectren Conservation Park
in 2002.

Family            Scientific Name

Caesalpiniaceae   Cercis canadenses
Cornaceae         Cornus amomum
Cornaceae         Cornus florida
Cornaceae         Cornus racemosa
Cornaceae         Nyssa sylvatica
Fagaceae          Quercus alba
Fagaceae          Quercus imbricaria
Fagaceae          Quercus macrocarpa
Fagaceae          Quercus michauxii
Fagaceae          Quercus palustres
Hamamelidaceae    Liquidambar styraciflua
Juglandaceae      Carya illinoinensis
Juglandaceae      Juglans nigra
Lauraceae         Lindera benzoin
Oleaceae          Fraxinus pennsylvanica
Plantanaceae      Plantanus occidentales
Rosaceae          Crataegus phaenopyruin
Rosaceae          Physocarpus opulifoius
Rubiaceae         Cephalanthus occidentales

Family            Common Name            Inds. Planted

Caesalpiniaceae   redbud                      2300
Cornaceae         silky dogwood               2000
Cornaceae         flowering dogwood           2300
Cornaceae         gray dogwood                2300
Cornaceae         black gum                   7000
Fagaceae          white oak                  20900
Fagaceae          shingle oak                 4600
Fagaceae          bur oak                    10000
Fagaceae          swamp chestnut oak          4500
Fagaceae          pin oak                    10000
Hamamelidaceae    sweetgum                   20000
Juglandaceae      pecan                      10000
Juglandaceae      black walnut               10000
Lauraceae         spicebush                   2300
Oleaceae          green ash                  10000
Plantanaceae      sycamore                   13000
Rosaceae          Washington hawthorne        2300
Rosaceae          ninebark                     700
Rubiaceae         buttonbush                  1900
                  Total trees planted       136100


APPENDIX 2:

Species list of flora present at Vectren Conservation Park (arranged alphabetically by family). Each species report includes the following information: (1) scientific name based on Gleason and Cronquist (1991), (2) common name, (3) origin (native or introduced), (4) wetland indicator category (U.S. Army Corps of Engineers 1996), and (5) coefficient of conservatism (C-value) for Indiana (Rothrock 2004). For the wetland indicator categories, OBL represents obligate wetland plants (with plants almost always occurring in wetlands (>99%)), FACW represents facultative wetland plants (with plants usually occurring in wetlands (67%-99%)), FAC represents facultative plants (with plants being equally likely to occur in wetlands or non-wetlands (34%-66%)), FACU represents facultative upland plants (with plants usually occurring in non-wetlands (67%-99%)), and UPL represents Obligate Upland plants (with plants almost always occurring in non-wetlands in our region (>99%))(Reed 1996). NI represents plants with insufficient information available to determine indicator status. Signs (+/-) represent discrimination within categories, with a positive sign representing a greater frequency in that habitat and a negative sign representing a lower frequency). C-values range from zero to 10, with lower values representing plants that are highly tolerant of disturbance and higher values representing plants that are restricted to high quality plant community remnants. In addition to the species listed only to the level of genus, eight species were identified only at higher taxonomic levels; four specimens belonged to three families--Brassicaceae, Poaceae, and Asteraceae, and four were not placed to any family. Many introduced plants were neither categorized for wetland category nor for C-values. Still, introduced species are often given C-values of zero.

Acanthaceae (Acanthus Family) Ruellia strepens L.: Wild petunia; native; FAC+; 4

Aceraceae (Maple Family) Acer negundo L.: Boxelder; native; FACW-; 1 Acer saccharinum L.: Silver maple; native; FACW; 1

Amaranthaceae (Amaranth Family) Amaranthus sp.

Apiaceae (Carrot Family)

Cryptotaenia canadensis (L.) DC.: Honewort; native; FAC; 3

Torilis arvensis (Huds.) Link: Field hedge-parsley; introduced, Europe

Apocynaceae (Dogbane Family)

Apocynum cannabinum L.: Indian hemp; native; FAC; 2

Araceae (Arum Family)

Arisaema dracontium (L.) Schott: Green dragon; native; FACW; 5

Aristolochiaceae (Birthwort Family) Aristolochia tomentosa Sims: Pipe vine; native; FAC; 7

Asclepiadaceae (Milkweed Family)

Ampelamus albidus (Nutt.) Britton: Bluevine; native;--; 1

Asclepias incarnata L.: Swamp milkweed; native; OBL; 4

Asclepias syriaca L.: Common milkweed; native; NI; 0

Asteraceae (Aster Family)

Achillea millefolium L.: Yarrow; native; FACU; 0

Ambrosia artemisiifolia L.: Common ragweed; native; FACU; 0

Ambrosia trifida L.: Giant ragweed; native; FAC+; 0 Aster praealtus Poir.: Veiny lined aster; native; FACW; 6

Aster pilosus Willd.: Heath aster; native; 0

Bidens comosa (A. Gray) Wiegand: Strawstem burmarigold; native;--; 1

Carduus nutans L.: Musk thistle; introduced, Europe

Conyza canadensis (L.) Cronquist: Horseweed; native; FAC-; 0

Erigeron annuus (L.) Pers.: Daisy fleabane; native; FAC-; 0

Erigeron philadelphicus L.: Philadelphia fleabane; native; FACW; 3

Eupatorium coelestinum L.: Mistflower; native; FACW; 2

Eupatorium serotinum Michx.: Late boneset; native; FAC+; 0

Iva annua L.: Rough marsh elder; native; FAC; 0

Lactuca serriola L.: Prickly lettuce; introduced, Europe; FAC; 0

Pyrrhopappus carolinianus (Walter) DC.: False dandelion; introduced; 2

Rudbeckia hirta L.: Black-eyed Susan; native; FACU; 2

Rudbeckia laciniata L.: Cutleaf coneflower; native; FACW; 3

Rudbeckia triloba L.: 3-lobed coneflower; native; FAC 3

Senecio glabellus Poir.: Butterweed; native; OBL; 0

Solidago canadensis L.: Common goldenrod; native; FACU; 0

Tragopogon dubius Scop.: Yellow salsify; introduced, Europe

Verbesina alternifolia (L.) Britton: Wingstem; native; FACW; 3

Vernonia gigantea (Walter) Trel.: Tall ironweed; native; FAC; 2

Bignoniaceae (Trumpet Creeper Family)

Campsis radicans (L.) Seem.: Trumpet creeper; native; FAC; 1

Catalpa speciosa Warder: Northern catalpa; native; FACU; 0

Brassicaceae (Mustard Family)

Capsella bursa-pastoris (L.) Medik.: Shepherd's purse; introduced, Europe; FAC-

Cardamine rhomboidea (Pers.) DC.: Springcress; native

Lepidium virginicum L.: Poor man's pepper; native; FACU-; 0

Rorippa palustris (L.) Besser: Common yellow cress; native; OBL; 2

Rorippa sessiliflora (Nutt.) Hitchc.: Southern yellow cress; native; OBL; 3

Caesalpiniaceae (Caesalpinia Family)

Cercis canadensis L.: Redbud; native; FACU; 3

Gleditsia triacanthos L.: Honey-locust; native; FAC; 1

Gymnoeladus dioicus Lam.: Kentucky coffee-tree; native; 3

Campanulaceae (Bellflower Family)

Campanula americana L.: Tall bellflower; native; FAC; 4

Caprifoliaceae (Honeysuckle Family) Symphoricarpos sp.

Caryophyllaceae (Pink Family) Dianthus armeria L.: Deptford pink; introduced, Europe; NI

Chenopodiaceae (Goosefoot Family) Chenopodium sp.

Convolvulaceae (Morning-glory Family)

Calystegia sepium (L.) R. Br.: Hedge bindweed; native; FAC; 4

Ipomoea hederacea Jacq.: Ivy-leaved morning glory; introduced; FAC Ipomoea pandurata (L.) G.Mey.: Wild potato; native; FACU; 3

Cornaceae (Dogwood Family)

Cornus drummondii C. A. Mey.: Rough-leaved dogwood; native; FAC; 2

Cucurbitaceae (Gourd Family)

Sicyos angulatus L.: Bur cucumber; native; FACW-; 3

Cuscutaceae (Dodder Family)

Cuscuta gronovii Willd. Common dodder; native; 2

Cyperaceae (Sedge Family)

Carex conjuncta Boott: Green-headed fox sedge; native; FACW; 6

Carex cristatella Britton: Crested sedge; native; FACW+; 3

Carex digitalis Willd.: Slender woodland sedge; native; UPL; 7

Carex grayi J. Carey: Globe sedge; native; FACW+; 5

Carex muehlenbergii Schkuhr ex Willd.: Muehlenberg's sedge; native;--; 5

Cyperus strigosus L.: False nutsedge; native; FACW; 0

Elaeagnaceae (Oleaster Family)

Elaeagnus angustifolia L.: Russian olive; introduced, native of Eurasia; FACU-Fabaceae (Pea Family)

Amorpha fruticosa L.: False indigo; native; OBL; 3 Desmodium paniculatum (L.) DC.: Panicled tickclover; native; FACU; 2

Lespedeza cuneata (Dum. Cours.) G. Don: Silky bushclover; introduced, Asia; UPL

Melilotus albus Medik: White sweet clover; introduced, Asia;

Melilotus officinalis (L.) Pall.: Yellow sweet clover; introduced, Eurasia; FACU

Trifolium campestre Schreb.: Low hop clover; introduced, Eurasia and N. Africa

Fagaceae (Beech Family)

Quercus macrocarpa Michx.: Bur oak; native; FAC-; 5

Geraniaceae (Geranium Family)

Geranium carolinianum L.: Carolina crane's bill; native;--; 2

Juglandaceae (Walnut Family)

Juglans cinerea L.: Butternut; native; FACU+; 5 Juglans nigra L.: Black walnut; native; FACU; 2 Lamiaceae (Mint Family)

Lamium amplexicaule L.: Henbit deadnettle; introduced, Eurasia and Africa

Teucrium canadense L.: American germander; native; FACW-; 3

Liliaceae (Lily Family)

Allium vineale L.: Field garlic; introduced, native of Europe; FACU

Malvaceae (Mallow Family)

Hibiscus laevis All.: Smooth rose mallow; native; OBL; 4 Sida spinosa L.: Prickly mallow; native; FACU

Menispermaceae (Moonseed Family)

Menispermum canadense L.: Common moonseed; native; FAC; 3

Mimosaceae (Mimosa Family)

Desmanthus illinoensis (Michx.) MacMill.: Bundle-flower; native; FACU; 3

Moraceae (Mulberry Family)

Maclura pomifera (Raf.) Schneid: Osage orange; native; FACU

Morus alba L.: White mulberry; introduced, Asia; FAC Morus rubra L.: Red mulberry; native; FAC-; 4

Oleaceae (Olive Family)

Forestiera acuminata (Michx.) Poir.: Swamp privet; native; OBL; 8

Fraxinus americana L.: White ash; native; FACU; 4

Fraxinus pennsylvanica Marshall: Green ash; native; FACW; 1

Onagraceae (Evening Primrose Family) Oenothera biennis L.: Common evening primrose; native; FACU; 0

Oenothera laciniata Hill: Ragged evening primrose; native; FACU; 2 Oenothera sp.

Oxalidaceae (Wood-sorrel Family)

Oxalis dillinii Jacq.: Southern yellow wood-sorrel; native; NI; 0

Phytolaccaceae (Pokeweed Family)

Phytolacca americana L.: American pokeweed; native; FAC-; 0

Plantaginaceae (Plantain Family)

Plantago rugelii Decne.: American plantain; native; FAC; 0

Platanaceae (Plane-tree Family)

Platanus occidentalis L.: American sycamore; native; FACW; 3

Poaceae (Grass Family)

Agrostis gigantea Roth: Redtop; introduced, Europe; FACW

Andropogon virginicus L.: Broomsedge; native; FAC-; 1

Arundinaria gigantea (Walter) Muhl.: Giant cane; native; NI, 5

Bromus racemosus L.: Bald brome; introduced, Europe

Chasmanthium latifolium (Michx.) H.O. Yates: Wild oats; native; FAC; 4

Dactylis glomerata L. Orchard grass; introduced, Europe; FACU

Echinochloa crusgalli (L.) P.Beauv.: Barnyard grass; introduced, Eurasia; FACW

Elymus canadensis L.: Canada wildrye; native; FAC-; 5

Elymus virginicus L.: Virginia wildrye; native; FACW-; 3

Festuca pratensis Huds.: Meadow fescue; introduced, Europe; FACUFestuca subverticillata (Pers.) E.B. Alexeev: Nodding fescue; native; FACU+; 4

Hordeumjubatum L.: Foxtail barley; native; FAC+ Hordeum pusillum Nutt.: Little barley; introduced, N. America; FAC; 0

Koeleria pyramidata (Lam.) P. Beauv.: Junegrass; native;--; 8

Lolium perenne L.: Perennial rye; introduced, Europe; FACU

Phleum pratense L.: Common timothy; introduced, Europe; FACU

Setaria viridis (L.) P. Beauv.: Green foxtail; introduced, Europe

Sorghum bicolor (L.) Moench: Sorghum; introduced, Africa; UPL

Sorghum halepense (L.) Pers.: Johnson grass; introduced, Europe and Africa; FACU

Polygonaceae (Smartweed Family)

Polygonum aviculare L.: Knotweed; native; FACPolygonum cuspidatum Siebold & Zucc: Japanese knotweed; introduced; Japan; FACU

Polygonum erectum L.: Erect knotweed; native; FACU; 0

Polygonum pennsylvanicum L.: Pinkweed; native; FACW+; 0

Polygonum persicaria L.: Spotted lady's thumb; introduced, Europe; FACW

Polygonum virginianum L.: Jumpseed; native; FAC; 3

Rurnex altissimus A. W. Wood: Pale dock; native; FACW-; 2

Rumex crispus L.: Curley dock; introduced, Europe; FAC+

Ranunculaceae (Buttercup Family)

Ranunculus micranthus Nutt.: Small-flowered crowfoot; native; FAC-; 4 Rosaceae (Rose Family)

Crataegus mollis (Torr. & Gray) Scheele: Downy hawthorn; native; FACW-; 2

Geum canadense Jacq.: White avens; native; FAC; 1 Potentilla norvegica L.: Rough cinquefoil, native; FAC; 0 Prunus sp.

Rubus sp. Rubiaceae (Madder Family)

Cephalanthus occidentalis L.: Common button-bush; native; OBL; 5

Salicaceae (Willow Family)

Populus deltoides Marshall: Cottonwood; native; FAC+; 1

Salix exigua Nutt.: Sandbar willow; native; FACW+; 1 Salix nigra Marshall: Black willow; native; OBL; 3

Saururaceae (Lizard's tail Family)

Saururus cernuus L.: Lizard's tail; native; OBL; 4

Scrophulariaceae (Figwort Family) Verbascum blattaria L.: Moth mullein; introduced, Eurasia; FACUMimulus alatus Alton: Sharpwing monkey-flower; native; OBL; 4

Smilacaceae (Catbriar Family)

Smilax herbacea L.: Smooth carrion flower; native; FAC; 4 Smilax sp.

Solanaceae (Nightshade Family)

Solanum carolinense L.: Horse-nettle; native; FACU-; 0

Ulmaceae (Elm Family)

Celtis laevigata L.: Southern hackberry; native; FACW; 7

Celtis occidentalis L.: Northern hackberry; native; FAC-; 3

Ulmus americana L.: American elm; native; FACW-; 3 Ulmus rubra Muhl.: Slippery elm; native; FAC; 3

Urticaceae (Nettle Family) Laportea canadensis (L.) Wedd.: Canada nettle; native; FACW; 2

Pilea pumila (L.) A. Gray: Clearweed; native; FACW; 2 Urtica dioica L. Stinging nettle; native; FACW-; 1

Valerianaceae (Valerian Family)

Valerianella radiata (L.) Dufr.: Beaked cornsalad; native; FAC; 1

Verbenaceae (Vervain Family)

Verbena urticifolia L.: White vervain; native; FAC+; 3

Phyla lanceolata (Michx.) Greene: Fogfruit; native; OBL; 2

Violaceae Viola sp.

Vitaceae (Grape Family)

Parthenocissus quinquefolia (L.) Planch.: Virginia creeper; native; FAC-; 2

Vitis aestivalis Michx.: Summer grape; native; FACU; 4

Vitis cinerea Engelm.: Sweet winter grape; native; FACW-; 4

Vitis riparia Michx.: Riverbank grape; native; FACW-; 1

Vitis vulpina L.: Winter grape; native; FACW-; 3

ACKNOWLEDGMENTS

We thank Vectren Corporation for their generous support and creative thinking, which provided the land on which the study was conducted. We appreciate their gift and support for restoration at VCP. Thanks to Allen Rose and Randy Kron for assistance at VCP, Jack Barner and UE's development office for their assistance, Dr. Alan Kaiser for assistance with historical land use information, Dr. Ann Powell for assistance with plant identification, and Dr. Dale Edwards and Kayla Stilger for their comments on earlier drafts of this manuscript. This work has been supported by a Merck/AAAS Undergraduate Science Research grant, a UExplore grant from the Undergraduate Research Committee at the University of Evansville, and a Foundation for Sustainability and Innovation to CGH. We thank an anonymous reviewer for helpful guidance.

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Jordon Lachowecki, Cris G. Hochwender *, Kristen Nolting, Abby Aldridge and Elizabeth Maurer: Department of Biology, University of Evansville, 1800 Lincoln Ave., Evansville, IN 47722

* Corresponding author. ch81@evansville.edu, Department of Biology, University of Evansville, 1800 Lincoln Ave., Evansville, IN 47722, 812-488-2005, 812-488-1039 (fax)

Table 1.--Species encountered in (A) meadow and (B) forest plots
at Vectren Conservation Park. Family, origin (native or
introduced), scientific and common  name, and number of stems are
given. A dash in the origin column indicates uncertain origin;
these species were not included in abundance calculations.

A. Meadow

Family           Origin           Scientific Name

Aceraceae        Native           Acer saccharinum
Anacardiaceae    Native           Toxicodendron radicans
Apiaceae         Introduced       Torilis arvensis
Asteraceae       Native           Ambrosia artemisifolia
Asteraceae       Native           Ambrosia trida
Asteraceae       Native           Aster praealtus
Asteraceae       --               Aster sp. 1
Asteraceae       Native           Cony--a canadensis
Asteraceae       Native           Erigeron annuus
Asteraceae       Native           Lactuca canadensis
Asteraceae       Introduced       Lactuca serriola
Asteraceae       Native           Pyrrhopappus carolinianus
Asteraceae       Native           Solidago canadensis
Asteraceae       Native           Taraxacum officinale
Brassicaceae     Introduced       Capsella burla pastoris
Convolvulaceae   Native           Calystegia sepium
Chenopodiaceae   --               Chenopodium sp.
Fabaceae         Introduced       Lespedeza cuneata
Fabaceae         Introduced       Medicago lupulina
Fabaceae         Introduced       Melilotus albus and Mehlotus
                                    officinalis
Fabaceae         Introduced       Trifolium campestre
Onagraceae       Native           Oenothera biennis
Oxalidaceae      --               Oxalis sp.
Polygonaceae     --               Polygonum sp.
Poaceae          Native           Andropogon virginicus
Poaceae          Native           Elymus virginicus
Poaceae          Introduced       Sorghum halepense
Ulmaceae         Native           Ulmus rubra and U. americana
Verbenaceae      Native           Verbena urticifolia

B. Forest

Family           Origin           Scientific Name

Anacardiaceae    Native           Toxicodendron radicans
Apiaceae         Native           Cryptotaenia canadensis
Asteraceae       Native           Ambrosia trifda
Asteraceae       --               Aster sp. 2
Asteraceae       --               Aster sp. 3
Asteraceae       Native           Solidago canadensis
Bigoniaceae      Native           Campis radicans
Convolvulaceae   Introduced       Ipomea hederacea
Cyperaceae       Native           Carex grayi
Menispermaceae   Native           Menispermum canadense
Oleaceae         Native           Fraxinus pennsylvania
Phytolaccaceae   Native           Phytolacca americana
Polygonaceae     Native           Polygonum virginianum
Poaceae          Native           Elymus virginicus
Poaceae          Native           Festuca subverticillata
Smilaceae        Native           Smilax herbacea
Smilaceae        --, but native   Smilax sp.
Solanaceae       Native           Solanum ptycanthum
Ulmaceae         Native           Celtis laevigata
Urticaceae       Native           Loportea canadensis
Urticaceae       Native           Pilea pumila
Vilaceae         --               Viola sp.
Vitaceae         Native           Parthenocissus quinquefolia

A. Meadow

Family           Common Name                    Plot I   Plot II

Aceraceae        Silver maple (sapling)            1         0
Anacardiaceae    Poison ivy                        1         0
Apiaceae         Field hedge parsley               0         1
Asteraceae       Common ragweed                    0         1
Asteraceae       Great ragweed                    40         6
Asteraceae       Veiny-line aster                870       328
Asteraceae       Unidentified long-leaf aster    145         3
Asteraceae       Horseweed                         0         5
Asteraceae       Daisy fleabane                    1         0
Asteraceae       Wild lettuce                      0        11
Asteraceae       Prickly lettuce                   0         0
Asteraceae       False dandelion                   0         1
Asteraceae       Common goldenrod                162       353
Asteraceae       Common dandelion                 11        41
Brassicaceae     Shepherd's purse                  0         1
Convolvulaceae   Hedge bindweed                    1         1
Chenopodiaceae   Lamb's quarters                   0         0
Fabaceae         Silky bushclover                  0        48
Fabaceae         Black medic                       1         0
Fabaceae         Sweet clover                      0      1159

Fabaceae         Low hop clover                    5        20
Onagraceae       Biennial evening primrose         1        40
Oxalidaceae      Woodsorrel                        1         0
Polygonaceae     Various smartweeds                0         0
Poaceae          Broom sedge                      22         0
Poaceae          Virginia wildrye                392        79
Poaceae          Johnson grass                    23       156
Ulmaceae         Elm (sapling)                     0         4
Verbenaceae      White vervain                     1         0

B. Forest

Family           Common Name                    Plot I   Plot II

Anacardiaceae    Poison ivy                       16        14
Apiaceae         Honewort                        690       245
Asteraceae       Great ragweed                     2         0
Asteraceae       Unidentified forest aster 1      42         0
Asteraceae       Unidentified forest aster 2      27         3
Asteraceae       Common goldenrod                  0         0
Bigoniaceae      Trumpet creeper                   0         2
Convolvulaceae   Ivy-leaved morning glory          2         0
Cyperaceae       Globe sedge                      22       175
Menispermaceae   Common moonseed                   0         0
Oleaceae         Green ash                         0         0
Phytolaccaceae   American pokeweed                26         0
Polygonaceae     Jumpseed                          1         5
Poaceae          Virginia wildrye                490       758
Poaceae          Nodding fescue                    0         0
Smilaceae        Smooth carrion-flower             1         1
Smilaceae        Unidentified smilax               3        13
Solanaceae       Eastern black nightshade          0         0
Ulmaceae         Sugarberry (sapling)              0         0
Urticaceae       Wood nettle                      97       112
Urticaceae       Clearweed                       317        14
Vilaceae         Unidentified violet               3        12
Vitaceae         Virginia creeper                 38        44

A. Meadow

Family           Plot III   Total

Aceraceae             0        1
Anacardiaceae        18       19
Apiaceae              0        1
Asteraceae           16       17
Asteraceae            4       50
Asteraceae           87     1285
Asteraceae            1      149
Asteraceae           92       97
Asteraceae            0        1
Asteraceae            2       13
Asteraceae            8        8
Asteraceae            0        1
Asteraceae           55      570
Asteraceae           85      137
Brassicaceae          0        1
Convolvulaceae        2        4
Chenopodiaceae      519      519
Fabaceae              0       48
Fabaceae           3234     3235
Fabaceae             70     1229

Fabaceae              0       25
Onagraceae          466      507
Oxalidaceae           0        I
Polygonaceae          8        8
Poaceae               0       22
Poaceae             155      626
Poaceae             274      453
Ulmaceae              0        4
Verbenaceae           3        4

B. Forest

Family           Plot III   Total

Anacardiaceae        17       47
Apiaceae              7      942
Asteraceae            0        2
Asteraceae           36       78
Asteraceae            0       30
Asteraceae            3        3
Bigoniaceae           1        3
Convolvulaceae        0        2
Cyperaceae          351      548
Menispermaceae        1        1
Oleaceae              1        1
Phytolaccaceae        0       26
Polygonaceae          0        6
Poaceae             627     1875
Poaceae               8        8
Smilaceae             1        3
Smilaceae             0       16
Solanaceae            4        4
Ulmaceae              5        5
Urticaceae            1      210
Urticaceae          342      673
Vilaceae              4       19
Vitaceae             59      141
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Author:Lachowecki, Jordon; Hochwender, Cris G.; Nolting, Kristen; Aldridge, Abby; Maurer, Elizabeth
Publication:Proceedings of the Indiana Academy of Science
Geographic Code:1U3IN
Date:Jul 20, 2012
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