Journal of the Torrey Botanical Society 132(2), 2005, pp. 323–354

Vascular plants of a Delaware River tidal freshwater wetland and adjacent terrestrial areas: Seed bank and vegetation comparisons of reference and constructed marshes and annotated species list1

Mary Allessio Leck2,3 Biology Department, Rider University, Lawrenceville, NJ 08648

Charles F. Leck Department of Ecology and Evolutionary Biology, Cook College, Rutgers University, New Brunswick, NJ 08901

LECK, M. A. (Dept. Biol., Rider Univ., Lawrenceville, NJ 08648) AND C. F. LECK (Dept. Ecol. & Evol. Biol., Cook College, Rutgers Univ., New Brunswick, NJ 08901). Vascular plants of a Delaware River tidal freshwater wetland and adjacent terrestrial areas: Seed bank and vegetation comparisons of reference and constructed marshes and annotated species list. J. Torrey Bot. Soc. 132: 323–354. 2005.—Constructed wetland soil samples contained many more species with densities Ͼ 100 seeds / m2 than reference marsh ones (102 vs. 28); constructed wetland densities for many species were considerably higher and richness (species / sample) were Ͼ 2ϫ greater than the reference marsh. Of the total 218 seed bank species, 60 were common to both and 34 occurred only in reference marsh samples. Numbers of cover species were similar, but composition differed. Invasive species, notably Lythrum salicaria, Phalaris arundinacea, and Phragmites australis, were more important in constructed wetland soil samples and vegetation. Non-native species comprised 9 % and 13 % of the reference marsh and constructed wetland seed bank species and 8 % and 12 % of the cover species, respectively. Variable dispersal patterns and in situ seed production contributed to these differences. Over the entire study area (wetlands and adjacent terrestrial areas) we recorded 875 species, representing 141 families, more than doubling the number observed in a 1988 report. The families having the most species were Asteraceae (103 species), Poaceae (100), and Cyperaceae (83). The largest genera were Carex (46) and Polygonum (19); seven other genera also had Ն 10 species. Nine species were pseudo-viviparous, producing plantlets on inflorescences. Non-native species comprised 27 % of the total flora. Overall there were 37 NJ rare / endangered species for the entire area; 11 occurred only in the constructed wetland with constructed wetland soil samples containing more than the ref- erence marsh samples (8 vs. 1). Available lists from 1824, 1887, and 1964/65 and five species known only from herbarium specimens indicate that several species have been extirpated since the 19th C. Anthropogenic influences continue to have impact on diversity. Key words: biodiversity, constructed wetland, floristic survey, reference marsh, tidal freshwater marsh, wetland seed bank.

The Delaware River tidal freshwater marsh In ton—Trenton—Bordentown or Trenton Marsh, New Jersey, USA, locally known as the Hamil- has been the focus of many vegetation studies beginning in the 1970s (e.g., Whigham and 1 Rider University provided financial and other sup- Simpson 1975, Simpson et al. 1983, Leck and port. 2 We could not have accomplished this survey with- out invaluable help from many including: Patrick Coo- formation about the constructed wetland plantings. ney, Robert Meyer, Bill Olson, William Standaert, and Herb Lord prepared the map. others who participated in Philadelphia Botanical Many students and friends have accompanied us Club, Torrey Botanical Society, and other field trips. and/or helped with various aspects of our field and Karl Anderson, David Fairbrothers, Ted Gordon, Linda greenhouse work. We especially note the assistance of Kelly, John Kuser, Janice Meyer, James Montgomery, Fay Josephson, Robin MacKinnon, Patricia Orban and David Snyder helped with troublesome specimens. Quinby, Donna Wilson, Jennifer Wilson and Marian We are especially indebted to A.E. Schuyler and An- Young. Robert Simpson provided helpful comments on thony Reznicek; Ernie Schuyler helped identify many an earlier draft of the manuscript. We appreciate the species when we were new to wetland plants, and as careful reading and helpful suggestions of two review- we continued our survey both he and Tony Reznicek ers. Colleen Dillon helped prepare the appendix in the helped with difficult graminoids. form it appears here; Elizabeth Faulkenstein and Rich- Patricia Tyson Stroud provided a copy of her hand- ard Deni helped with previous versions. To these and written notes taken from Charles Lucien Bonaparte’s others we extend our sincere thanks. letters. Lucy Aiello lent several of Charles Conrad Ab- 3 Author for correspondence, E-mail: [email protected] bott’s books and Ned Gilmore provided copies of ar- Received for publication August 19, 2004, and in ticles by C.C. Abbott. Michael Kaminsky provided in- revised form December 14, 2004.

323 324 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

FIG. 1. Map of Hamilton—Trenton—Bordentown Marsh, including adjacent terrestrial habitats included in this survey. Numbered locations are: 1 Beaver Point, 2 Spring Lake trails, 3 Osprey Point, 4 Sunny Pond, 5 little mitigation site, 6 Sturgeon Pond (east and west), 7 landfill, 8 Rowan Lake, 9 Watson Woods, 10 site of Abbott’s homestead, 11 reference / pre-existing marsh, 12 Hamilton sewage disposal plant, 13 Bordentown City water works, 14 Northern Community Park (Bordentown Township), 15 Public Service Gas & Electric Co., 16 gas and oil tanks, 17 Duck Island constructed wetland, 18 abandoned brick factory, 19 lagoon, 20 Bordentown Bluffs, 21 Duck Creek, 22 Conrail (Camden and Amboy) Railroad, 23 Delaware & Raritan Canal, 24 Delaware & Raritan Canal State Park, 25 Point Breeze, 26 railroad bridge, 27 Black Creek.

Simpson 1995, Leck 2003). In 1988 we sum- make comparisons of both seed bank and veg- marized qualitative and quantitative data on dis- etation of a reference marsh area and a con- tribution and abundance of plant species and structed wetland and evaluate the impact of hu- provided descriptions of location, physiography, man-generated landscape disturbance. Such a sediments, and geology as well as land use his- comparison, particularly based on long-term tory (Leck et al. 1988). At that time we recorded studies, has not been made previously for this 426 species. wetland and there are few studies comparing ref- The goals of this report are to provide a status erence and constructed (or restored) wetlands summary of vegetation at two levels. First, we that include seed bank data. Baldwin and De- 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 325

Rico (1999) is an exception. Second, we provide which has been the location of several seed bank an annotated and updated view of plant species studies (e.g., Leck and Graveline 1979, Leck diversity that can provide insights regarding the and Simpson 1995). The entire study area, in- quality of wetland and adjacent terrestrial plant cluding terrestrial areas, will be referred to sim- communities, as well as a basis for encouraging ply as the ‘Marsh’. management activities and developing educa- tional uses (Delaware & Raritan Greenway, SITES FOR SEED BANK AND VEGETATION 1999). These could also provide a baseline for STUDIES. The reference marsh was a small por- ϳ future floristic studies (Robinson et al. 1994). tion ( 10 ha) of the 505 ha pre-existing marsh. Its location was chosen for accessibility and be- cause it had representative tidal marsh plant Survey Area. OVERVIEW. The Marsh (Fig. 1) is located 3.5 km south of Trenton, New Jersey. communities (see Whigham and Simpson 1975). Ͻ It lies between industrialized land on Duck Is- It is fed by two shallow ( 2 m) tidal channels ϳ ϳ land and urbanized land, associated with growth and is 1 km from Crosswicks Creek and 0.5 of Trenton, Hamilton Township, Bordentown km from Watson Creek. The sediments are fine City, Bordentown Township, and other commu- silt and clay that contain as much as 36 % or- nities. Wetlands are bounded by the Delaware ganic material (Simpson et al. 1983, Leck et al. River on the west and bordered by bluffs to the 1988). north, east (and southeast), and south. The sur- In contrast, the constructed wetland has vey area also included the Crosswicks Creek coarse sediments derived from river cobble- floodplain to NJ Route 130 near Yardville, the stones, sand, and gravel present at the time of Delaware River shore from Bordentown to the construction. It is linked to the Delaware River ϳ Trenton boat launch on Lamberton Road, Duck by four short ( 20 m) inlets lined with riprap. Creek, and terrestrial areas within the wetland The sampling locations, except that on Elatine and along the bluffs. These areas are contiguous Island, were accessible by foot. (See Leck with the wetland itself and comprise an entity 2003). that is relatively unaffected by the dense urban HABITAT TYPES. Wetlands. Wetland com- development around it. This survey included munities are varied and include 18 riverine and species from a variety of habitats (see below). palustrine communities identified by Whigham The designated area covers nearly 1215 ha, of and Simpson (1975) (see also Leck et al. 1988). which about 550 ha are wetlands. These are found in stream channel, stream bank, Since the previous study, two new wetlands high marsh, and tidal pond locations. To these have been constructed along the Delaware River can be added constructed wetlands, non-tidal on Duck Island to compensate for wetland lost ponds, such as Spring Lake, and beaver im- during construction of the Trenton Highway poundments in former pond-like areas. Beaver, Complex and for river edge lost during construc- whose activity was first noted in 1996, have had tion of NJ-129 along the Delaware River in substantial impact raising water level in the Trenton, NJ. The former, a 38.9 ha site com- pond-like areas near Spring Lake by more than pleted in 1994, is the largest New Jersey De- 0.5 m and impeding tidal water flow except dur- partment of Transportation wetland mitigation ing flood tides. Occasionally, flood tides destroy project in New Jersey. It has 32.3 ha of wet- the dams, causing partial and temporary draw- lands, of which about 70% are tidal. There are downs. ϳ 2 km of channels and eight islands (from northeast to south, island names are D.O.T., Ab- Tidal Rivers and Channels. A number of bott, Dorothy Cross, Elatine, Kalm, Tide Pool, tidal freshwater rivers define this wetland; these Lenape, and Tiny Islands). For construction and include the Delaware River and Crosswicks, other details see Leck (2003). In 2002, a second Watson, and Duck Creeks, which have a 2.3 m 1 ha constructed wetland was completed. tidal range. Much of the eastern bank of the Del- For clarity, the marsh areas will be designated aware River along the edge of the Marsh has a as follows: of the approximately 550 ha of wet- cobble bottom where certain plant species able lands, 38.9 ha on Duck Is. comprise the ‘con- to tolerate winter scouring by water and ice, e.g., structed wetland’ (Fig. 1, #17); the remainder Aster puniceus, Helenium autumnale, Lythrum will be referred to as the ‘pre-existing marsh’. salicaria, Vernonia noveboracensis, and in few- Near the east edge of the pre-existing marsh is er locations Justicia americana, may be found located the ‘reference marsh’ (Fig. 1, #11), near the high tide line. (For species authorities 326 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132 see Appendix 1). Below these and also in the vey area. Marshes may be found at the edges of intertidal zone, occur Polygonum punctatum and ponds, along tidal channels, or other poorly Nuphar lutea. Rare species such as Bidens bi- drained areas. They may be dominated by pe- dentoides may be found along the upper inter- rennial species, such as Leersia oryzoides, Pon- tidal cobbles and Sagittaria subulata in lower tederia cordata,orTypha spp. In 2004 Phalaris intertidal mudflats. Except for the west bank of arundinacea was prominent along the stream Duck Creek (near its mouth), the banks of other channel levee in the reference marsh. Along tid- rivers are of fine sediments. Where water move- al channels annual species may also seasonally ment is rapid no macrophytes grow. However, dominate the vegetation (Simpson et al. 1983). submergent species such as Ceratophyllum de- These include Amaranthus cannabinus, Ambro- mersum, Elodea nuttallii, Myriophyllum spica- sia trifida, Polygonum punctatum, and Zizania tum, Vallisneria americana, and Zannichellia aquatica. Back from the channel edge the high palustris can occur in sizeable beds in Duck marsh may be dominated by Bidens laevis, Cus- Creek, Watson Creek, and the Delaware River. cuta gronovii, Impatiens capensis, and Polygo- In shallow channels Nuphar lutea and Zizania num arifolium, interspersed with perennials, aquatica may clog the waterways in summer. such as Acorus calamus, Peltandra virginica, and Schoenoplectus fluviatilis. In tidal marsh ar- Ponds. Sturgeon Pond, Spring Lake, and eas, with marked seasonal changes from mud or Rowan Lake (tidal), as well as areas impounded litter in March, to a carpet of seedlings in April, by the highway construction of I-195, I-295, and to vegetation reaching 3 m tall in August, the NJ-129 interchange, and the pond in the Duck prodigious growth places these tidal freshwater Island constructed wetland are to some degree marshes among the most productive ecosystems the result of human alteration. For example, in the world (Tiner 1985). Spring Lake, also known as White City Lake, was constructed in the late 19th century as part Shrub Forest Swamps. Tidal and nontidal of an amusement park. Since 1996, beaver have shrub forest wetlands that are characterized by created additional impoundments. The ponds are woody species occur at the edges of marshes of different ages and depths and have distinctive adjacent to terrestrial areas. The thickets of Al- characteristics. Both Spring Lake and Sturgeon nus incana, Alnus serrulata, Cephalanthus oc- Pond show the effects of hydrological succes- cidentalis, Cornus amomum, Ilex verticillata, sion. Spring Lake has extensive coverage of Nu- and/orViburnum dentatum, with Acer rubrum phar lutea and a fringe of emergents, such as and Fraxinus pennsylvanica, occur on hum- Lythrum salicaria, Peltandra virginica, Ponted- mocks surrounded by marsh. Some shrub forest eria cordata, and Typha latifolia. Sturgeon areas support showy species such as Aster pun- Pond, which was divided in half by highway iceus, Cardamine bulbosa, Chelone glabra, Lil- construction in the 1980s, in contrast, has a ium canadense, Lysimachia ciliata, and Rosa broad zone of floating and emergent species palustris, but where there is a dense canopy with Ludwigia peploides, Nuphar lutea, Peltan- these showy species are lacking. dra virginica, and Polygonum amphibium grad- A variant of this habitat are alder swamps, ing into Hibiscus moscheutos, shrubs (e.g., Vi- with Carex stricta tussocks, other Cyperaceae, burnum dentatum and Alnus spp.), and trees Acer rubrum, and Alnus spp., that are located at (e.g., Acer rubrum). The tidal Rowan Lake is the bottom of the bluffs near the C.C. Abbott dominated by Nuphar lutea, which is emergent homestead and along the Crosswicks Creek throughout the tidal cycle. Ponds contain sub- floodplain near Yardville. Only scattered sedge merged and floating macrophytes that do not oc- tussocks have been observed in other areas. This cur in other wetland types. These include Nym- vegetation type may have been historically more phaea odorata (mostly extirpated by 2004). extensive; it was documented in a photograph Ludwigia peploides, which is infrequent in New published in the 19th C (Abbott 1898). The Ab- Jersey (Hough 1983), and Proserpinaca palus- bott swamp site may be compromised both by tris. Utricularia radiata and Lemnaceae, includ- isolation caused by highway construction and by ing New Jersey rare / endangered species Wolf- storm drain runoff. fiella gladiata and Lemna valdiviana (Snyder Constructed Wetlands. The constructed wet- 1997), have been locally abundant. land, because of its proximity to the Delaware Marshes. Examples of both tidal and nontid- River, openness, and coarse soil that is different al herbaceous marshes are found within the sur- from the fine silty soils found elsewhere, has 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 327 provided habitat for unusual species, including mum, as well as native grasses, including Schi- Senna hebecarpa and Elsholtzia ciliata.Inad- zachyrium scoparium and Sorgastrum nutans, dition, Elatine americana and Heteranthera can be found. However, once the canopy devel- multiflora occurred in shallow ‘tide pool’ de- ops these are lost. Second growth woods can pressions. Woody species, including Betula ni- also be observed on Duck Island north of and gra, Platanus occidentalis, Rubus spp., Salix around the perimeter of the constructed wetland, spp. and Vitis spp., quickly became established. as well as along the Delaware & Raritan Canal It has also provided transitory habitat for many and railroad tracks. There Robinia pseudoacacia other species. Low stature species, such as Lin- is common and the rare Ptelea trifoliata may dernia dubia, Ludwigia palustris, and Eleochar- occur in open areas is acicularis were replaced by taller species in- Except for destruction caused by a tornado cluding Aster puniceus, Impatiens capensis, during late June 1996, relatively undisturbed Lythrum salicaria, Phragmites australis, and woods occur on the bluffs along Crosswicks others. See Leck (2003). Creek from Point Breeze to the Bordentown The little mitigation site has also provided Bluffs. These are dominated by Quercus alba, habitat for opportunistic species, but all have Q. montana, and Q. rubra, with Liriodendron been found in other areas. tulipifera and Nyssa sylvatica. A few trees of Pinus rigida and Tsuga canadensis also occur Terrestrial Habitats. Forests vary in soil there. Unique for this part of New Jersey are type and plant communities they support. West dense thickets of Kalmia latifolia and Rhodo- of Spring Lake is a low-lying wooded area dom- dendron maximum that dominate the shrub layer. inated by Acer rubrum and Liquidambar styra- ciflua. Beaver activity since 1996 has made the Disturbed Areas. Construction and mainte- area wetter, killing trees whose roots could not nance activities along roadways, the railroad, tolerate prolonged inundation. Nine species of sewage treatment plants, and other buildings, as ferns, including Botrychium virginianum and well as places where flotsam is deposited by Woodwardia areolata, have been found along tides, offer obviously altered and disturbed en- trails. vironments. These occur on a variety of soil Wet woods may also be found on the flood types and are places to look for plants that grow plain of Crosswicks Creek up stream of the in open, often ephemeral habitats. Among the Route 206 bridge near Yardville. Tree species more interesting species found in such places there include Acer negundo, Acer saccharinum, have been Coreopsis lanceolata, Heterotheca Fraxinus spp., Nyssa sylvatica, Platanus occi- subaxillaris, Paulownia tomentosa, and Vicia te- dentalis, and Salix spp. A small stand of mature trasperma. Chamaecyparis thyoides also occurs in this area. Highway construction, which began in 1984, Vines of Vitis spp., Menispermum canadense, caused considerable disturbance in some areas. Parthenocissus quinquefolia, and Toxicoden- Such open places provided a transitory habitat dron radicans festoon the trees along the stream for a variety of species. For example, during a and riverbanks. During spring, wildflowers, such Torrey Botanical Society / Philadelphia Botani- as Claytonia virginica, Erythronium american- cal Club trip in September 1990, 12 Polygonum um, Floerkea proserpinacoides, Mertensia vir- species were observed along the road edge (I- ginica, and Viola spp., are abundant. Symplo- 195 cloverleaf) at the east edge of the marsh. carpus foetidus is locally common in seepage Several mustards in disturbed areas, including areas. Camelina microcarpa, Diplotaxis tenuifolia, and Drier woodlands may be found along the tops Sisymbrium altissimum, are infrequent in NJ and of the bluffs, on bluff slopes and on the islands have not been previously recorded for Mercer within the pre-existing marsh. They are domi- County (Hough 1983). nated by Carya spp., Fagus grandifolia, Liriod- endron tulipifera, Quercus spp., and other hard- Methods. COMPARISONS OF REFERENCE AND wood trees. These woods support a variety of CONSTRUCTED WETLANDS. Comparisons are gen- wild flowers, such as the spring-blooming Di- erally based on examination of published stud- centra cucullaria, Podophyllum peltatum, Smi- ies: Leck (2003) for the Duck Island constructed lacina racemosa, and Uvularia sessilifolia.In wetland and Leck and Graveline (1979), Parker open sandy areas in summer, Chamaecrista nic- and Leck (1985), Leck et al. (1988) and Leck titans, Lespedeza spp., and Trichostema dicoto- and Simpson (1987a, b, 1994, 1995) for the ref- 328 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132 erence marsh. These document the size and Desmodium cuspidatum (likely Desmodium can- composition of the soil seed bank, as well as adense), Euphorbia chamissonis (actually Cha- composition and % cover of the field vegetation. maesyce nutans), and Pycnanthemum muticum In addition, vegetation changes of the construct- (likely Pycnanthemum tenuifolium). ed wetland were documented on one of the is- Generally, nomenclature and authorities fol- lands (Elatine) in the following manner. During low Rhoads and Klein (1993) and, if a species late summer (14 September 1995, 30 August is not recorded for Pennsylvania, we used Glea- 1996, 29 August 1997, 28 August 1998, 21 Au- son and Cronquist (1991). For Kyllinga we used gust 1999, 20 / 28 August 2000, 28 August Bryson, et al. (1997). We have changed earlier 2001, 13 September 2002, 30 August 2003, 7 listings (Leck et al. 1988) to reflect the more September 2004), a 165 m transect was estab- recent taxonomy (e.g., Alliaria officinalis to A. lished that extended 30 m into upland areas and petiolata). To determine modern synonymies for 105 m across a tidal marsh saddle with a 30 m several species given in Bonaparte (1824) and perpendicular extension reaching to the closest Abbott (1886 1887), we used Gray (1887); in tidal channel. At 15 m points along the transect some cases clues came from common names 5 m diameter circular plots were marked. Within that are still in use. For New Jersey rare or en- each plot both common (a subjective estimate of dangered species status designation we used species having the highest densities) and species Snyder (1997). present were recorded. Results and Discussion. Our previous study SPECIES SURVEY. The species list (Appendix (Leck et al. 1988) provided a hint of the diver- 1) is based on plants germinating in soil sam- sity of vascular plant species found in the Marsh ples, occurring in field vegetation plots, and/or with 426 species recorded; here we increase the found during extensive field surveys conducted number to 875 (Appendix 1). The increase can both on foot and by canoe. It also includes spe- be attributed to: (1) continuing and expanded cies listed in Leck et al. (1988), and other pub- field exploration, but also (2) the considerable lications, dating to vegetation studies undertaken habitat disturbance related to highway construc- by Whigham and Simpson (1975), unpublished tion between 1984–1994 (some of this effect data, as well as species observed by J. Arsenault, was included in the 1988 report); (3) construc- R. Blicharz, and A. E. Schuyler (including spec- tion of mitigation wetlands on Duck Island, 38.9 imens in the A. E. Schuyler Herbarium at the ha in 1994 and 1 ha in 2002, that provided new Academy of Natural Sciences of Philadelphia). habitat for many species and (4) location of Although available 19th and 20th century lists these constructed wetlands adjacent to the Del- are incomplete, they provide glimpses of the aware River that provided a large and diverse past flora (Bonaparte 1824, Abbott, 1886, 1887, seed resource (Leck 2003); and finally (5) in- 1892, 1897, 1898, Shoemaker 1964, 1965). Such creasing the survey area by including the Bor- species are noted on Appendix 1 or listed in Ap- dentown Bluffs, Northern Community Park, pendix 2. The Appendix 2 species from Bona- Delaware & Raritan Canal State Park lands, and parte, Abbott, and Shoemaker are not included other areas, not previously explored, along the in tallies. floodplains of Crosswicks Creek near Yardville, Appendix 3 lists species planted at the con- Duck Creek, and the Delaware River shore. In structed wetland. Five, not part of the regional addition, our comparisons of the reference flora (Hough 1983) nor observed elsewhere in marsh and constructed wetland provide a view the Marsh, are also not included in tallies. of how disturbance in a wetland system with Herbarium specimens for 690 species, main- ample seed dispersal can promote diversity (Ta- tained at Rider University, have allowed us to bles 1, 2, 3). check many of our original identifications (Leck We will first consider comparisons between et al. 1988). Several listings from the 1988 study the reference marsh and the constructed wetland. that were given only to genus (e.g., Lilium sp.), Then we will examine the entire flora and pro- based on vegetative plants, are not included. Of vide some historical notes. the species listed by Leck et al. (1988), the fol- lowing are not included here because their ac- COMPARISONS OF THE REFERENCE MARSH AND curate identification was questioned: Berberis CONSTRUCTED WETLAND. Numerous compari- vulgaris (actually Berberis thunbergii), Celas- sons can be made between the constructed wet- trus scandens (actually Celastrus orbiculatus), land and reference marsh, based on seed banks 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 329 and vegetation data, that illustrate the high spe- deposition of seed-ladden flotsam there (Leck cies diversity and colonization potential of tidal 2003). freshwater marshes, as well as the importance of Species diversity along hydrologic gradients rare or endangered and non-native species. The had different patterns along transects from seed bank of the constructed wetland had 177 stream channels to the high marsh. The refer- taxa germinating in soil samples (over 5 years); ence marsh had somewhat higher numbers along of these 92 contributed densities Ͼ 100 / m2 (Ta- the stream channel compared with high marsh ble 1; Leck 2003). In contrast, the reference locations (Parker and Leck 1985, Leck and marsh samples contained only 96 species (over Simpson 1994), while in the constructed wet- 15 years) and 28 taxa had densities exceeding land, numbers were lowest at the channel edge 100/m2. Moreover, of the 101 seed bank spe- (Leck 2003). In the constructed wetland, espe- cies contributing more than 100 seeds / m2, only cially in the first 2–3 years, there was little per- 18 were common to both; nine others were sistent plant material near the channel edge to found only in the reference marsh and 83 only trap seeds, and the tides deposited the seeds well in constructed wetland samples. A total of 218 away from the edge. In the reference marsh seed plant species occurred in seed bank sam- plants growing on the channel levee likely re- ples; of these 124 occurred only in constructed duced water flow and, thus, trapped seeds. wetland samples and 34 only in reference marsh Both constructed wetland and reference marsh ones. In addition, many more Asteraceae, Cy- studies showed that there were typically fewer peraceae, and Juncaceae species occurred in species in field plots than in seed bank samples. constructed wetland than in reference marsh For example, reference marsh field plots con- samples (Tables 1, 2; Appendix 1). These distri- tained 24 species, with 18 species contributing butions suggest dispersal limitations of many to cover, compared to 96 species in soil samples (Leck et al. 1988); the constructed wetland had seed species. 92 species in field plots, of which 72 contributed In contrast to seed plants, more spore-bearing to cover, compared to 177 in soil samples (Leck species, seven fern and one horsetail species, 2003). These differences between field and grew from reference marsh samples, but only greenhouse data may be due to optimal germi- Onoclea sensibilis from constructed wetland nating conditions in the greenhouse for more ones. The lack of other ferns in the constructed species compared to the field, but also it is also wetland samples may be due to limited avail- possible that field conditions, such as a coating ability spores, but may also be due to use of tap of mud and high growth rates of other species, instead of distilled water for watering and/or to were unsuitable for establishment especially of colonization by a greenhouse moss contaminant small seedlings. that could have influenced germination and Overall, the numbers of species contributing growth of spore species. to cover were similar with 66 species occurring Many more constructed wetland species had in the reference marsh plots (all studies) and 72 high seed bank densities with certain species species in the constructed wetland. Composition, greatly exceeding the maximum total seed bank however, was not similar. In the reference marsh, ϭ values for any reference marsh site (max. depending on location relative to the tidal chan- Ϯ Ϯ 64,300 8,218 ( SE), Leck and Simpson nel, dominants were Acorus calamus, Bidens 1987a) (Table 1). Richness values in the con- laevis, Impatiens capensis, Peltandra virginica, ϫ structed wetland were more than 2 greater Polygonum arifolium, and Typha latifolia (all Ϯ Ϯ ϭ (22.9 1.0 to 27.5 1.7 species / sample; n species having Ͼ 10 % cover; Leck and Simp- 18) than those of the reference marsh (5.1 Ϯ 0.4 son 1987a, 1995), as well as Amaranthus can- to 10.8 Ϯ 0.6) (0–2 cm; March samples; Leck nabinus, Nuphar lutea, Polygonum punctatum, and Simpson 1987a). However, during early suc- and Sagittaria latifolia (Ͼ 5% relative density cession in the constructed wetland, patterns var- or cover; Parker and Leck 1985). Most of these ied with location; richness values of edge areas reference marsh species were absent as cover did not decrease (Leck 2003), but did on Elatine species during early succession from the con- Island (Fig. 2). This pattern of species richness structed wetland; instead Aster puniceus, Juncus decline during early succession is similar to that effusus, Lindernia dubia, Ludwigia palustris, of abandoned agricultural fields (Leck and Leck Lythrum salicaria, Mimulus spp., and Pentho- 1998). Continued high species numbers along rum sedoides were important (Table, 1; Leck edges may be due to water current patterns and 2003). Furthermore, the constructed wetland 330 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Table 1. Seed bank species contributing Ͼ 100/m2 in reference (RM) and constructed wetland (CW) studies*. Data are maximum mean densities (Ϯ SE). For the constructed wetland seed bank frequencies are also given and are based on four years, three sites and three locations (n ϭ 36). (Overall, total species numbers were: RM ϭ 95, CW ϭ 183; RM ϩ CW ϭ 218, with 34 in RM only and 60 in both).

Density Species (Mean Ϯ SE) Freq. (%) REFERENCE MARSH (28 taxa) Amaranthus cannabinus 3,675 Ϯ 1,044 Ambrosia trifida 119 Ϯ 42 Bidens laevis 1,690 Ϯ 554 Boehmeria cylindrica Ͼ100 Callitriche heterophylla 915 Ϯ 401 Cardamine pensylvanica 500 Ϯ 397 Carex spp. (3) 280 Ϯ 136 Cuscuta gronovii 4,830 Ϯ 2,390 Dulichium arundinaceum 1,040 Ϯ 560 Eupatorium dubium 180 Ϯ 106 Gratiola neglecta Ͼ100 Impatiens capensis 2,080 Ϯ 315 Juncus effusus 1,280 Ϯ 731 Leersia oryzoides 190 Ϯ 158 Mikania scandens 8,360 Ϯ 1,636 Nuphar lutea 120 Peltandra virginica 141 Ϯ 80 Phalaris arundinacea 3,433 Ϯ 3,433 Pilea pumila 6,350 Ϯ 1,524 Poa trivialis 2,540 Ϯ 1,944 Polygonum arifolium 2,415 Ϯ 524 Polygonum punctatum 1,475 Ϯ 591 Polygonum sagittatum 140 Ranunculus sceleratus 320 Ϯ 287 Sagittaria latifolia 425 Ϯ 115 Sium suave 125 Ϯ 73 Typha latifolia 53,000 Ϯ 7,445 Zizania aquatica 163 Ϯ 138 CONSTRUCTED WETLAND (100 taxa) Agrostis perennans 183 Ϯ 183 2.8 Alisma plantago-aquatica 100 Ϯ 100 5.6 Amaranthus cannabinus 985 Ϯ 602 44.4 Artemisia vulgaris 200 Ϯ 113 25 Arthraxon hispidus 933 Ϯ 856 66.7 Aster pilosus 1,633 Ϯ 848 63.1 Aster puniceus 10,300 Ϯ 2,471 50 Aster spp. 1,833 Ϯ 164 22.2 Bidens connata 150 Ϯ 50 19.4 Bidens frondosa 550 Ϯ 263 44.4 Bidens laevis 4,617 Ϯ 2,308 61.1 Boehmeria cylindrica 3,243 Ϯ 737 69.4 Brassicaceae 185 Ϯ 153 27.8 Carex (I) 617 Ϯ 368 66.7 Carex (II) 514 Ϯ 481 30.6 Carex (III) 414 Ϯ 351 50 Carex frankii 150 Ϯ 150 11.1 Carex lurida 133 Ϯ 133 5.6 Carex scoparia 1,350 Ϯ 1,105 22.2 Carex spp. 666 Ϯ 523 19.4 Carex stipata 1,350 Ϯ 1,350 8.3 Carex stricta 135 Ϯ 135 2.8 Carex tribuloides 350 Ϯ 212 19.4 Carex vulpinoidea 116 Ϯ 116 22.2 Cephalanthus occidentalis 100 Ϯ 37 13.9 Chenopodium ambrosioides 1,686 Ϯ 386 75 Cicuta bulbifera 100 Ϯ 63 8.3 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 331

Table 1. Continued.

Density Species (Mean Ϯ SE) Freq. (%) Cuscuta gronovii 1,883 Ϯ 324 30.6 Cyperus bipartitus 7,250 Ϯ 2,879 58.3 Cyperus diandrus 267 Ϯ 158 13.9 Cyperus erythrorhizos 1,971 Ϯ 963 50 Cyperus lupulinus 166 Ϯ 148 16.7 Cyperus flavescens 616 Ϯ 617 13.9 Cyperus odoratus 2,617 Ϯ 863 2.8 Cyperus strigosus 3,486 Ϯ 637 80.6 Cyperus spp. 850 Ϯ 540 38.9 Digitaria filiformis 128 Ϯ 89 8.3 Digitaria ischaemum 133 Ϯ 49 25 Echinochloa crusgalli 3,017 Ϯ 1,878 77.7 Eclipta prostata 542 Ϯ 76 55.6 Eleocharis acicularis 11,200 Ϯ 10,000 16.7 Eleocharis erythropoda 133 Ϯ 133 8.3 Eleocharis obtusa 4,583 ϩ 2,120 77.8 Eleocharis spp. 166 Ϯ 166 16.7 Erigeron annuus 1,183 Ϯ 596 11.1 Eupatorium perfoliatum 3,183 Ϯ 1,571 33.3 Eupatorium serotinum 150 Ϯ 62 47.2 Euthamia graminifolia 283 Ϯ 125 27.8 Gratiola neglecta 17,316 Ϯ 4,222 75 Helenium autumnale 983 Ϯ 887 47.2 Hypericum mutilum 2,400 Ϯ 1,875 69.4 Impatiens capensis 583 Ϯ 214 38.8 Juncus acuminatus 14,120 Ϯ 6,651 86.1 Juncus effusus 39,967 Ϯ 20,311 75 Juncus tenuis 12,400 ϩ 8,290 75 Juncus spp. 7,017 ϩ 4,081 50 Leersia oryzoides 2,550 Ϯ 1,175 77.8 Lemna minor 752 Ϯ 752 8.3 Lindernia dubia 172,214 Ϯ 23,235 97.2 Ludwigia alternifolia 117 Ϯ 83 13.9 Ludwigia palustris 258,610 Ϯ 34,080 97.2 Lycopus americanus 1,050 Ϯ 665 30.6 Lycopus europaeus 750 Ϯ 243 27.8 Lycopus virginicus 150 Ϯ 72 19.4 Lycopus spp. 6,016 Ϯ 4,177 58.3 Lythrum salicaria 207,900 Ϯ 30,760 88.9 Microstegium vimineum 666 Ϯ 539 50 Mikania scandens 4,757 Ϯ 3,102 16.7 Mimulus ringens (ϩalatus) 4,767 Ϯ 2,560 92.7 Mollugo verticillata 550 Ϯ 347 38.9 Oenothera biennis 250 Ϯ 56 36.1 Panicum dichotomiflorum 2,333 Ϯ 2,095 75 Panicum spp. 267 Ϯ 193 19.4 Penthorum sedoides 6,150 Ϯ 1,819 86.1 Phalaris arundinacea 1,133 Ϯ 1,133 47.2 Phragmites australis 133 Ϯ 133 27.8 Pilea pumila 14,900 Ϯ 4,849 72.2 Platanus occidentalis 117 Ϯ 65 16.7 Poa trivialis 133 Ϯ 114 8.3 Polygonum arifolium 128 Ϯ 57 (ϩ) Polygonum caespitosum 883 Ϯ 289 86.1 Polygonum hydropiper 1,733 Ϯ 870 61.1 Polygonum hydropiperoides 883 Ϯ 472 66.7 Polygonum lapathifolium 1,167 Ϯ 632 77.8 Polygonum pensylvanicum 883 Ϯ 313 83.3 Polygonum persicaria 117 Ϯ 117 19.4 332 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Table 1. Continued.

Density Species (Mean Ϯ SE) Freq. (%) Polygonum punctatum 8,933 Ϯ 2,453 83.3 Polygonum setaceum 220 Ϯ 120 30.6 Polygonum spp. 1,500 Ϯ 664 50 Potentilla norvegica 3,117 Ϯ 1,154 80.6 Sagittaria latifolia 2,300 Ϯ 803 47.2 Schoenoplectus smithii 300 Ϯ 300 27.8 Scirpus cyperinus 173 Ϯ 173 2.8 Scutellaria epilobiifolia 200 Ϯ 113 19.4 Scutellaria lateriflora 133 Ϯ 67 27.8 Sium suave 167 Ϯ 80 11.1 Solidago canadensis 650 Ϯ 422 44.1 Solidago rugosa 100 Ϯ 68 38.9 Typha spp. 7,100 Ϯ 7,078 50 Verbascum thapsus 285 Ϯ 184 25 Verbena hastata 283 Ϯ 245 22.2 Veronica peregrina 1,050 Ϯ 637 38.8 * Data are from Leck and Graveline (1979), Parker and Leck (1985), Leck and Simpson (1987a, 1994, 1995) for the reference marsh, and Leck (2003) for the constructed wetland, and unpublished data. cover species, except for Gratiola neglecta, of the reference marsh are associated with tran- were found only occasionally or not at all in the sience, where a sizable component of seed bank reference seed bank samples over 15 years lasts Ͻ 1 yr in the soil, while small seeds, char- (Leck and Simpson 1987a, 1994, 1995, Leck et acteristic of the constructed wetland, are persis- al. 1988). tent and last Ͼ 1 yr (Leck and Simpson 1987a, Certain reference marsh dominants occurred 1995; Leck and Brock 2000). in the constructed wetland, but varied in their Particular groups, such as rare or endangered colonization potential. Two, Bidens laevis and and non-native species, differed between wet- Polygonum punctatum, were common during the lands. During the early colonization period, first year (Leck 2003). With time, however, Im- more rare or endangered species occurred in the patiens capensis and Cuscuta gronovii became constructed wetland; soil samples of the refer- more important in the vegetation; so that after ence marsh contained only one rare species four years Impatiens capensis occurred in 93 % compared with eight in constructed wetland of upland edge plots, with cover averaging 33.5 samples (Table 3). Some, such as Isoetes ripar- Ϯ 6.4 % (n ϭ 27), and in 62 % of the plots (n ia, were scarce and transitory, with only a few ϭ 13) on Elatine Island. Cuscuta gronovii oc- plants found during a single growing season. curred in 26 % of the cover plots (contributing Non-native species comprised 9 % of the seed to cover only in 7 % of the plots with maximum bank species in the reference marsh and 13 % cover of 5 %; n ϭ 27) and in 38 % of plots on in the constructed wetland, and 8 and 12 %, re- Elatine Island. spectively, of the cover species. These percent- In addition to these density and diversity com- ages are low compared to other wetland studies parisons, vegetation structure and morphology (e.g., Magee et al. 1999) reflecting the need for differed. Colonization of the constructed wet- wetland specialist species that are able to toler- land was by many low-growing species. Taller ate, for example, variable tidal inundation re- species quickly became established contributing gimes and rapid growth of other species (see Az- to a layered community (Leck 2003). The ref- nar et al. 2003). Of the invasive non-native spe- erence marsh did not demonstrate layering be- cies considered to be problems in wetlands, cause nearly all species were capable of tall and/ Lythrum salicaria, Phalaris arundinacea, and or viney growth (Leck and Simpson 1995). Field Phragmites australis were more important in production of pseudo-viviparous plantlets oc- both the seed bank and vegetation of the con- curred only in the constructed wetland. Also, the structed wetland (Table 1; Appendix 1) and did colonizing dominant species of the constructed not appear in the reference marsh vegetation wetland (1995–1999) had smaller seed sizes plots (Leck and Simpson 1994, 1995) although (0.01–3.2 mg; n ϭ 8) than those of the reference Phalaris arundinacea did establish along the marsh (0.01 to 530 mg; n ϭ 19). The large seeds channel levee (MAL pers. observ.). In the con- 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 333 0 7.7 e highest 71.4 33.3 70 53 69.9 66.7 63.3 37.9 88 50 80 29.4 75 26.6 33.3 60 22.2 77.8 55.5 25 25 ercents of CW f % total 141 443 875 236 es species, and species 0 5 2 5 4 1 4 6 2 7 5 2 2 spp. 70 53 58 22 19 11 22 11 16 12 CW f 339 22 55 105 260 0 4 1 4 7 2 3 5 0 1 1 0 3 1 1 3 2 2 0 spp. 29 23 33 13 CW sb 138 0 2 1 6 0 2 2 3 2 3 0 0 4 2 2 2 3 2 1 3 2 spp. 68 13 13 RM sb 103 311 580 181 % 0 0 0 2.4 0 0 0 0 27.2 37 57.6 30 27.6 40 59 40 41.2 62.5 20 25 40 12.5 37.5 of total 3 6 7 1 0 0 0 2 9 8 8 7 3 0 3 4 0 0 0 1 3 28 37 19 10 13 10 spp. 168 Nonnative 0 0 0 4.9 1 3.6 0 0 3.8 0 5 0 6.7 6.3 0 0 0 0 0 0 % 10 13.3 22.2 of total 7 0 14 21 0 2 5 1 3 0 4 0 1 0 1 0 1 1 0 0 1 0 2 0 0 0 0 spp. 22 Special 9 9 9 8 8 8 7 6 83 33 30 29 25 22 20 17 16 15 13 12 10 spp. Total 103 100 592 4 5 7 0 & Equisetophyta Pteridophyta Pinophyta Magnoliopsida Liliopsida Total Lycopodiophyta Table 2. Vascular plant summaries for the Hamilton–Trenton–Bordentown Marsh. A. Phyla summaries. B. Species distribution among families having th richness (see Appendix 1).distributions Shown for are: seed familyfamily banks species totals (sb) totals, are species of also having both given. special the designation reference (e.g., marsh rare or (RM) endangered; and Snyder constructed 1997), wetland non-nativ (CW) and for field cover (f) of the constructed wetland. The p Totals 875 37 4.2 236 27 117 177 465 53.1 A. Families Genera Species Non-native species B. FAMILY Asteraceae Poaceae Cyperaceae Fabaceae Lamiaceae Rosaceae Polygonaceae Brassicaceae Scrophulariaceae Liliaceae Caryophyllaceae Salicaceae Fagaceae Apiaceae Ranunulaceae Dryopteridaceae Juncaceae Rubiaceae Betulaceae Caprifoliaceae Ericaceae Onagraceae Lemnaceae Subtotals 334 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Table 3. New Jersey rare or endangered species (Snyder 1997) recorded for the Hamilton–Trenton–Borden- town Marsh. Global and state rankings* are given. Species are designated as occurring in the reference marsh, constructed wetland, entire study area (Marsh), or as a herbarium record. Species located nearby study plots are ‘(ϩ)’. Species designated as ‘T’ occurred in terrestrial locations. Herbarium species are located in the A.E. Schuyler Herbarium of the Academy of Natural Sciences of Philadelphia. (From Leck et al. 1988, Leck 2003, and unpublished).

Reference marsh Constructed wetland Herbarium Species Seed bank Seed bank Vegetation Marsh record only Agastache nepetoides (G5, S2)* T Agastache scrophulariifolia (G4, S2) T Andropogon ternarius (G5, S2) T Aster longifolius (G5, S1, E) ϩ Bidens bidentoides (G3, S2, E) ϩ (ϩ) ϩ Cacalia atriplicifolia (G4-5, S1, E) T Carex frankii (G5, S3) ϩ (ϩ)T ϩ Carex sterilis (G4, S2) ϩ (ϩ) Cuscuta polygonorum (G5, S2) (ϩ) Cyperus retrofractus (G5, SH, E) T Elatine americana (G4, S2) (ϩ) Eriocaulon parkeri (G3, S2) ϩ Eupatorium altissimum (G5, S2) T Eupatorium aromaticum (G4-5, S1) T Euphorbia corollata (G5, S2) T Heteranthera multiflora (G4, S2) ϩ?(ϩ) ϩ Isoetes riparia (G4, S3) (ϩ) Juncus brevicaudatus (G5, S2) (ϩ) Juncus torreyi (G5, S1) (ϩ) Lemna valdiviana (G5, S1, E) ϩ Lycopus rubellus (G5, S2) ϩ (ϩ) Mimulus alatus (G5, S3) ϩ? (ϩ) ϩ Myriphyllum heterophylum (G5, S2) ϩ Nelumbo lutea (G4, S1, E) ϩ Polygonum setaceum (G5, S2) ϩ (ϩ) Populus heterophylla (G5, S2) ϩ ϩ Ptelea trifoliata (G5, S1, E) T T Quercus michauxii (G5, S3) ϩ Ranunculus longirostris (G5, S2) ϩ Sagittaria subulata (G4, S2) (ϩ) ϩ Schoenoplectus smithii (G5?, S2) ϩ (ϩ) Scirpus atrocinctus (G5, S1) (ϩ) Silene caroliniana (G5, S3) T Tradescantia ohiensis (G5, S2) T Utricularia gibba (G5, S3) ϩ Utricularia intermedia G5, S3) ϩ Wolffiella gladiata (G?, S1, E) ϩ Totals: 37 1 6 (8) 17 21 5 * Ranking (Snyder 1997): Global (G1–5); G1 ϭ critically imperiled globally to G5 ϭ demonstrably secure globally although possibly rare at the periphery of range; State (S1–5); S1 ϭ critically imperiled in New Jersey to S5 ϭ demonstrably secure in the state under present conditions. SH ϭ of historical occurrence in New Jersey, but no extant populations known. E ϭ endangered, prospects for survival in New Jersey are low and the species is in immediate danger. structed wetland, both on upland edge sites (Lis- sects (channel edge, midpoint, and upland edge cko unpublished) and along the study transect locations), vegetation frequencies were 0, 0, and on Elatine Island, Lythrum salicaria did not re- 33 %, respectively; after 10 years (2004) they duce species richness of the vegetation and, in were 0, 44, and 67% (n ϭ 9; Leck 2003 and fact, Lythrum salicaria itself declined after three unpublished data). In some locations, however, years (Fig. 2). Phragmites australis, as it contin- stands are not monospecific and Phragmites aus- ues to expand like it has in other wetlands (Rice tralis co-occurs with other species, such as Im- et al. 2000), may have the greatest and continu- patiens capensis and Mikania scandens. ing impact. After five years (in 1999) along tran- Population dynamics of these constructed 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 335

1995), cannot penetrate consolidated, compacted sediment (Garbisch 1994). In addition, wetland species are known to have variable germination responses to soil texture (Keddy and Constabel 1986, Weiher et al. 1996) and to sedimentation (Ditmar and Neely 1999). Another kind of comparison when evaluating the success of replacement wetlands, is whether constructed and reference marshes are equiva- lent. Studies from other locations have shown variable success (e.g., Bishel-Machung et al. 1996, Goldman-Carter and McCallie 1996, Gal- atowitsch and van der Valk 1996, Wilson and Mitsch 1996, Whigham 1999, Zedler and Cal- laway 1999, Klotzli and Grootjans 2001). Also, FIG. 2. Species richness on Elatine Island and oc- currences of invasive species. Data are numbers of although vegetation structure and seed banks common (bars) or present (black diamonds) species may be different from those of reference wet- (mean Ϯ SE) per plot. Also shown are the number of lands, constructed wetlands can provide com- plots in which Lythrum salicaria was common (open munity level information about seed banks and circles) and all plots in which Phragmites australis vegetation processes, including seed dispersal was found (triangles). (n ϭ 13; plots 5 m radius). patterns (Cappers 1993, Cain et al. 2000). They can also be used to assess vegetation structure wetland invasive species differed. Lythrum sal- and dynamics of constructed / restored versus icaria waxed and waned, but continued to con- reference wetlands (Baldwin and DeRico 1999, tribute to the vegetation (Fig 2). Overall, Phrag- Ehrenfeld 2000). Here, in the short time the con- mites australis, as noted above, appears to be structed wetland was monitored, the species-rich increasing although data from Elatine Is. (Fig. nature of tidal freshwater marshes (Simpson et 2) do not appear to indicate this. In contrast, al. 1983) was achieved, but species composition Phalaris arundinacea, present mainly at upland and structure were not similar (Table 1; Leck edges, was not found along study transects in 2003). 2004 (unpublished data). Thus, although the constructed wetland and Many factors contribute to the differences be- reference marsh did not have similar seed bank tween the reference marsh and the constructed densities, seed bank and vegetation species rich- wetland. These include reduced dispersal, relat- ness, or species composition, these comparisons ed to the relative isolation of the reference provide several insights. First, anthropogenic marsh, which receives water from two small me- manipulations and associated habitat diversifi- andering tidal channels connected to Crosswicks cation provided establishment habitats for many and Watson Creeks. For the constructed wetland species, including rare ones not found elsewhere the Delaware River (Fig. 1) was nearby and was in the Marsh. Second, colonizing (early succes- the source of flotsam that was Ͼ 10 cm deep in sional) species of the constructed wetland had many places and that contained large numbers high levels of in situ seed production, resulting of diverse seeds (MAL pers. observ.). In addi- in much higher densities and species richness tion to dispersal, conditions for establishment, values than those found in the reference marsh. although not fully understood, are involved. It is Finally, although numbers of non-native (and in- apparent that many species exploited the open vasive) species were a relatively small compo- niches associated with early successional habi- nent of the seed bank and vegetation for both tats following dispersal into the constructed wet- sites (8–13 %), their importance in the construct- land (Tables 1, 3; Fig.1). Such species were not ed wetland seed bank and vegetation was far able to tolerate increased competition as the veg- greater. etation developed and species of low stature were replaced by taller ones (Leck 2003). SPECIES SURVEY. To date (September 2004), Sediment differences may also contribute to we have recorded 875 species, including 148 establishment and seed bank patterns. For ex- (16.9 %) woody and 236 (27 %) alien species ample, roots of Peltandra virginica, one of the (Appendix 1). Thirty-seven (4.2 %) are worthy reference marsh dominants (Leck and Simpson of note as species that require conservation con- 336 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132 cern in New Jersey; of these, eight (0.9 %) are already present in 1994 (Leck 2003). Of the spe- designated as endangered (Table 3; Snyder cies listed for planting, we did not observe Zi- 1997). Parenthetically, 268 of these species were zania aquatica seedlings in 1995, but it was well not recorded for Mercer County by Hough established by 1998. Woody species appeared to (1983). The families having the greatest num- have fared the worst, generally because other bers of species are presented in Table 2. Overall, species, such as Robinia pseudoacacia, estab- 141 families were recorded; 11 were ferns and lished and grew quickly, overtopping them. fern allies, 3 Pinophyta, and 125 Magnoliophyta, Planted Acer rubrum and Sambucus canadensis the latter comprised of 103 Magnoliopsida and in some locations died or were stunted and chlo- 22 Liliopsida families. In the constructed wet- rotic, apparently due to improper location. Ter- land, species of certain families (Table 2 B), e.g., restrial areas, especially on islands, could be too Polygonaceae (88 %), Scrophulariaceae (80 %), dry in summer and too wet during winter flood- and Juncaceae (77.8 %), exceeded 70 % of the ing, reducing establishment. Planted macro- total species for that family. In contrast, Faga- phytes persisted at least until 1998, when after ceae (7.6 %) and Ericaceae (0 %) were not im- four years it was often not possible to distin- portant at the constructed wetland. In a recent guish plantings from the surrounding vegetation. evaluation of 70 geographically diverse seed The 875 species that are reported here for the bank studies, Leck and Schutz (2005) found the Marsh reflect the temporal and spatial diversity highest Cyperaceae diversity in constructed wet- of habitats. The occurrence of 37 rare or endan- land samples (Leck 2003). Eight genera provid- gered species, 11 of which occurred only in the ed Ͼ 10 species (Carex 46, Polygonum 19, Cy- constructed wetland (Table 3), also indicates the perus 13, Eupatorium 13, Panicum 12, Bidens importance of having varied habitats, including 11, Quercus 11, Salix 10). Eleven were errant or those with low fertility and low competition. planted ephemerals (end Appendix 1). The diverse habitats, briefly described above, Few species were documented from soil seed may be predictable, e.g., the reference marsh, bank samples (Leck et al. 1988, Leck 2003) that with regular tides and no seasonal drying (Leck were not also found in the field. These included: and Brock 2000) or unpredictable, e.g., sand Cyperus squarrosus, Drosera rotundifolia, Hed- bars and the constructed wetland, where sub- eoma puglegioides, Plantago major, Polygonum strate or quality of establishment niches change. careyi, Salix bebbiana, Salix hybrid, and Wood- Among habitats, there are also dynamic and var- wardia virginica. iable stresses (Table 4), with physical stresses Examination of herbarium specimens revealed being more important along stream channels and nine species, Cyperus bipartitus, C. flavescens, biotic stresses on the high marsh (Parker and Eleusine indica, Juncus acuminatus, Poa trivi- Leck 1985). Along tidal channels the scouring alis, Scirpus atrovirens, Scirpus georgianus, that occurred during winter removed all standing Scirpus polyphyllus, and Schoenoplectus purshi- dead plant material. In the constructed wetland anus, that produced pseudo-viviparous plantlets tidal deposition of wrack, coming from the near- on inflorescences. Except for Cyperus bipartitus by Delaware River, caused death of plants and and Poa trivialis, all were collected from the created open patches for colonization. Wrack de- field. position was not a noticeable feature of other In addition to those that dispersed into the marsh areas. site, 37 species were planted as part of the land- Factors contributing to high diversity include scaping provided for the Duck Island construct- effective dispersal, maintenance of ‘normal’ lev- ed wetland (NJDOT 1996) (Appendix 3). Plant- els of disturbance that involve, e.g., plant re- ing certainly helped initiate colonization. There placement (annuals), ice impacts in winter, sed- were at least two instances, Pontederia cordata iment deposition, etc., and changes in commu- and Sagittaria latifolia, where established plant- nities due, for example, to population dynamics ed plants provided a seed source for new seed- (Leck and Simpson 1995). Also, as noted above, lings (pers. observ.); only Sagittaria latifolia wetland creation and associated disturbance pro- was found in seed bank samples (Appendix 1). vided niches for many species. Reduction in di- However, contributions of the planted species to versity would come from inability of specialized the seed bank and vegetation were small because species to tolerate the changes. In some cases, so many other species grew there (Leck 2003). invasives may themselves be the disturbance, re- In fact, Peltandra virginica, Pontederia cordata, ducing diversity. While Lythrum salicaria, was and Sagittaria latifolia, likely from seeds, were not observed to reduce diversity in the con- 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 337

Table 4. Non-anthropogenic and anthropogenic stresses and main effects on tidal freshwater wetland vege- tation.

Stresses Main effects Non-anthropogenic Disturbance (litter/wrack, storms, animals) seed dispersal, seed rain, germination, recruitment Tide period & range (hydrology) germination, recruitment Biotic competition ⌬ recruitment, establishment Anthropogenic Climate change TЊ—germination, establishment Sea level rise communities change position along gradient; some com- munities lost locally Nutrients Ͼ growth, Ͼ biotic stress, Ͼ rate of germination (some species) Pollutants (e.g., pH: pesticides, oil, metals) recruitment, growth; indirect: herbivores, pollinators Invasive plant species Ͻ diversity, ⌬ seed rain patterns, availability of seeds Habitat fragmentation dispersal & seed rain patterns Sedimentation germination, recruitment, growth

structed wetland (Fig. 1; Liscko unpublished structed wetland was its substantial contribution data), expansion of Phragmites australis elimi- to species diversity (Tables 1–3, Appendix 1). nated species of low stature. However, extirpation of populations elsewhere could limit influx of propagules. Looking Ahead. Successional changes will continue to occur at the constructed wetland for SOME HISTORICAL NOTES. Glimpses of the for- some time. It is also anticipated that future mer botanical landscape are provided by Charles changes to the entire wetland complex will Lucien Bonaparte (1824), by Dr. Charles Conrad largely be anthropogenically induced. Orson et Abbott who wrote extensively about the area al. (1992) illustrate the considerable and con- around his home in the 19th century (e.g., Abbott tinuing human influence on marsh development 1887), and by Lois Shoemaker (1964, 1965). In as sea level rise and high sedimentation rates, addition to his own observations, Abbott pro- related to land use practices, support sediment vides a summary of those by Peter Kalm who levels conducive to tidal marsh communities. At in 1749 wrote about the plants he encountered the present time in the constructed wetland, sed- as he traveled by boat from Philadelphia to Tren- iment is accumulating in channels where water ton and then to New York City by stage. Kalm flow is slow. In addition, climate change, pol- reported that the forests near Trenton contained lutants (salts and other materials from highways an ‘‘abundance of chestnut-trees’’, and tulip- and storm drains), establishment of exotics and trees had replaced other species that had been invasives, management decisions, and habitat abundant during the previous century. He also fragmentation will affect wetland and terrestrial observed that, communities. Table 4 indicates some of the main ‘‘the beaver-tree (Magnolia virginiana) effects of various stresses on wetland plants. grows in the swamps. It is now (June 1st) These impacts are often subtle, but may be dra- in flower, and the fragrancy of its blossoms matic. has so perfumed the air that one could en- While species diversity, as represented by the joy it before one approached the swamps; total number of species found at the Marsh, in- and this fine smell likewise showed that a dicates high quality of ecosystems, watershed beaver-tree was near us, though we did not level management decisions are being made that happen to see it’’ (p. 147). may have significant (negative?) impact on the Marsh. Following storms, for example, there is Abbott (1887) concurred with Kalm, finding continuing high sediment load in Crosswicks evidence in local documents indicating that Creek related to watershed development. This magnolias had been more common two centuries study should serve as a baseline against which before. Today sweet bay magnolia is infrequent, to evaluate future anthropogenic impacts as well but there is a sizable tree at the north edge of as natural (e.g., successional) changes. Abbott’s swamp. Regarding the other tree spe- One of the most notable features of the con- cies mentioned, chestnuts are now infrequent 338 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132 due to chestnut blight. Tulip trees, in contrast, manipulation on plant species diversity in a and as noted by Kalm and Abbott, are still com- tidal freshwater system. mon and several large trees may be found along 3. The constructed wetland and little mitiga- the bluffs at Northern Community Park (Fig. 1). tion site contributed many species not (yet) Some of Abbott’s other observations include found elsewhere. his standing ‘‘in the midst of many acres’’ of 4. Many species were transitory in the con- bluets (Hedyotis caerulea), at a density of 16 per structed wetland where microhabitats square inch (Abbott 1887). He also wrote ‘‘In changed quickly during succession. an old woodland I once traced a fence by the 5. Large areas, including terrestrial habitats, long line of cypripediums in bloom, which were are critical to estimating and preserving spe- thriving in the mould of decayed fence-rails’ cies richness; several species occurred in a (Abbott 1898; p. 196). Golden club (Orontium single location. aquaticum) and lotus (Nelumbo lutea) were the 6. The constructed wetland, at least in its first two rarest plants (Abbott 1898). Similarly, we years, provided habitat for numerous spe- have observed dense patches, but not acres, of cies, including a number of rare ones that bluets in the Watson Woods picnic area and favor open, disturbed habitats. However, as know of only about a dozen golden club plants succession proceeds and as Phragmites aus- on the banks of Crosswicks Creek, perhaps in tralis expands its coverage, we predict open the same location where Abbott observed this habitat and species of low stature will be species. The lotus is no longer extant, but there further reduced. is a locally obtained specimen at the Academy 7. Continuing disturbance is key to providing of Natural Sciences of Philadelphia. Today lady habitat for colonizing species, species of slippers (Cypripedium acaule) appear to be low stature, and rare species, and for main- scarce and we have seen only five single and taining diversity. widely scattered non flowering plants although 8. Water is an effective dispersal agent for Bordentown residents reported having seen large many species, but many other wetland spe- numbers in flower during the past 30 years (Jan cies appear to have restricted distribution. Bisco Werner, pers. com.). 9. Historical records, however incomplete, Shoemaker (1964, 1965) also reported a num- provide clues to environmental change. ber of species for Roebling Memorial Park and 10. We expect that anthropogenic influences Charles Abbott’s homestead, called Three will continue to modify habitats and affect Beeches. For the latter, she noted honeysuckle, plant species establishment. We hope that the three beeches for which the property was management and open space decisions that named, a solitary white pine, as well as myrtle, are made for the Delaware River and Cros- daylilies, cultivated strawberries, hearts-ease swicks Creek watersheds will favor sustain- (Viola tricolor L.), and apple trees near the foun- able ecosystems in the study area. dation of the Abbott house cellar hole. The available lists of Bonaparte, Abbott, and Literature Cited Shoemaker and herbarium records indicate a loss or decline of a number of species (Appendix ABBOTT, C. C. 1886. Upland and meadow. Harper & 2, Table 3). For example, Nelumbo lutea and Brothers, New York, NY. Saxifraga virginiensis are no longer present, and ABBOTT, C. C. 1887. Wasteland wanderings. Harper & Aquilegia canadensis is found only on the rocks Brothers, New York, NY. ABBOTT, C. C. 1892. Recent rambles. J.B. Lippincott of the grotto at Point Breeze. Co., Philadelphia, PA. ABBOTT, C. C. 1897. The freedom of the fields. J.B. Conclusions. Lippincott Co., Philadelphia, PA. 1. We have doubled the number of species re- ABBOTT, C. C. 1898. Travels in a tree-top. J.B. Lippin- cott Co., Philadelphia, PA. ported in our 1988 inventory, suggesting AZNAR, J.-C., A. DERVIEUX, AND P. G RILLAS. 2003. As- that many still await discovery. It is our sociation between aquatic vegetation and landscape hope that the marshlands and surrounding indicators of human pressure. Wetlands 23: 149– terrestrial habitats will continue to inspire 160. botanical exploration. BALDWIN,A.H.AND E. F. DERICO. 1999. The seed bank of a restored tidal freshwater wetland in 2. Comparisons of reference and constructed Washington, D.C. Urban Ecosystems 3: 5–20. wetlands have contributed insights regard- BISHEL-MACHUNG, L., R. P. BROOKS,S.S.YATES, AND ing the positive impact of human landscape K. L. HOOVER. 1996. Soil properties of reference 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 339

wetlands and wetland creation projects in Pennsyl- LECK, M.A. AND C.F. LECK. 1998. A ten-year seed bank vania. Wetlands 16: 532–541. study of old field succession in central New Jersey. BONAPARTE, C.-L. 1824. Collection of Papers. Museum J. Torrey Bot. Soc. 125: 11–32. National d’Histoire Naturelle (Jardin des Plantes). LECK, M.A., AND W. S CHU¨ TZ. Regeneration of Cyper- Paris. (Notes provided by P. T. Stroud, March 2004; aceae, with particular reference to seed ecology and see also Stroud, P. T. 2001. The emperor of nature. seed banks. Persp. Pl. Ecol. Evol. System. University of Pennsylvania Press, Philadelphia, LECK,M.A.AND R. L. SIMPSON. 1987a. Seed bank of PA). a freshwater tidal wetland: turnover and relation- BRYSON, C. T., R. CARTER,L.B.MCCARTY, AND F. H . ship to vegetation change. Amer. J. Bot. 74: 360– YELVERTON. 1997. Kyllinga, a genus of neglected 370. weeds in the continental United States. Weed Tech. LECK,M.A.AND R. L. SIMPSON. 1987b. The spore 11: 838–842. bank of a Delaware River tidal freshwater wetland. CAIN, M. L., B. G. MILLIGAN, AND A. E. STRAND. 2000. Bull. Torrey Bot. Club 114: 1–7. Long-distance seed dispersal in plant populations. LECK,M.A.AND R. L. SIMPSON. 1994. Tidal fresh- Amer. J. Bot. 87: 1217–1227. water zonation: seed and seedling dynamics. CAPPERS, R. T. J. 1993. Seed dispersal by water: a con- Aquatic Bot. 47: 61–75. tribution to the interpretation of seed assemblages. LECK,M.A.AND R. L. SIMPSON. 1995. Ten-year seed Veg. Hist. Archaeobotany 2: 173–186. bank and vegetation dynamics of a tidal freshwater DELAWARE &RARITAN GREENWAY. 1999. Hamilton / marsh. Amer. J. Bot. 82: 1547–1557. Trenton Marsh management plan. (1 Preservation LECK, M. A., R. L. SIMPSON,D.F.WHIGHAM, AND C. Place, Princeton, NJ 08540). F. L ECK. 1988. Plants of the Hamilton Marshes: a DITTMAR,L.A.AND R. K. NEELY. 1999. Wetland seed Delaware River freshwater tidal wetland. Bartonia bank response to sedimentation varying in loading 54: 1–17. rate and texture. Wetlands 19: 341–351. MAGEE, T. K., T. L. ERNST,M.E.KENTULA, AND K. A. EHRENFELD, J. G. 2000. Defining the limits of resto- DWIRE. 1999. Floristic comparison of freshwater ration: the need for realistic goals. Restoration wetlands in an urbanizing environment. Wetlands Ecol. 8: 2–9. 19: 517–534. GALATOWITSCH,S.M.AND A. G. VAN DER VALK. 1996. NJDOT. 1996. Duck Island wetlands project—Trenton Characteristics of recently restored wetlands in the Complex I (195, I295, NJ 29, NJ129). Fact sheet prairie pothole region. Wetlands 16: 75–83. (distributed June). New Jersey Department of GARBISCH, E. 1994. The results of muskrat feeding on Transportation, Trenton, NJ. cattails in a tidal freshwater wetland. Wetland J. 6: ORSON, R. A., R. L. SIMPSON, AND R. E. GOOD. 1992. 14–15. The paleoecological development of a late Holo- GLEASON,H.A.AND A. CRONQUIST. 1991. Manual of cene, tidal freshwater marsh of the upper Delaware vascular plants of Northeastern United States and River estuary. Estuaries 15: 130–149. adjacent Canada. New York Botanical Garden, New York, NY. PARKER,V.T.AND M. A. LECK. 1985. Relationships of seed banks to plant distribution patterns in a fresh- GOLDMAN-CARTER,J.AND G. MCCALLIE. 1996. An en- vironmentalist’s perspective–time for a reality water tidal wetland. Amer. J. Bot. 72: 161–174. check, p. 88–108. In L. L. Marsh, D. R. Porter, and RHOADS,A.F.AND W. M. KLEIN,JR. 1993. The vas- D. A. Salvesen, (eds.), Mitigation banking theory cular flora of Pennsylvania, annotated checklist and and practice. Island Press, Washington, D.C. atlas. American Philosophical Society, Philadel- GRAY, A. 1887. Gray’s lessons in botany. The elements phia, PA. of botany for beginners and for schools. American RICE, D., J. ROOTH, AND J. C. STEVENSON. 2000. Col- Book Co., New York, NY. onization and expansion of Phragmites australis in HOUGH, M. Y. 1983. New Jersey wild plants. Harmony upper Chesapeake Bay tidal marshes. Wetlands 20: Press. Harmony, NJ. 280–299. KEDDY,P.A.AND P. C ONSTABEL. 1986. Germination of ROBINSON, G.R., M.E. YURLINA, AND S.N. HANDEL. ten shoreline plants in relation to seed size, particle 1994. A century of change in the Staten Island flo- size, and water level: an experimental study. J. ra: ecological correlates of species losses and in- Ecol. 74: 133–141. vasions. Bull. Torrey Bot. Club 121: 119–129. KLOTZLI,F.W.U.AND A. P. GROOTJANS. 2001. Resto- SHOEMAKER, L.M. 1964. Wild flowers along New Jer- ration of natural and semi-natural wetland systems sey highways and byways. Bull. 9, p. 1–25. New in Central Europe: progress and predictability of Jersey State Museum, Trenton, NJ. developments. Restoration Ecol. 9: 209–219. SHOEMAKER, L.M. 1965. Shrubs and vines of New Jer- LECK, M. A. 2003. Seed bank and vegetation devel- sey. Bull. 10, p. 1–54, New Jersey State Museum, opment in a created tidal freshwater wetland on the Trenton, NJ. Delaware River, Trenton, New Jersey, USA. Wet- SIMPSON, R. L., R. E. GOOD,M.A.LECK, AND D. F. lands 23: 310–343. WHIGHAM. 1983. The ecology of freshwater tidal LECK,M.A.AND M. A. BROCK. 2000. Ecological and wetlands. BioScience 33: 255–259. evolutionary trends in wetlands: evidence from SNYDER, D. B. 1997. Special plants of New Jersey. seeds and seed banks in New South Wales, Austra- Office of Natural Lands Management, Division of lia and New Jersey, USA. Plant Species Biol. 15: Parks and Forestry, New Jersey Department of En- 97–112. (Corrigendum 16: 183–184). vironmental Protection, Trenton, NJ. LECK, M.A. AND K.J. GRAVELINE. 1979. The seed bank TINER, R.W. 1985. Wetlands of New Jersey. US Fish of a freshwater tidal marsh. Amer. J. Bot. 66: 1006– and Wildlife Service, Region 5. Newton Corner, 1015. MA. 340 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

WEIHER, E., I. C. WISHEU,P.A.KEDDY, AND D. R. J. ment. Science of the Total Environment 240: 31– MOORE. 1996. Establishment, persistence, and man- 40. agement implications of experimental wetland WHIGHAM,D.F.AND R. L. SIMPSON. 1975. Ecological plant communities. Wetlands 16: 208–218. studies of the Hamilton Marshes. Progress Report WILSON,R.F.AND W. J. MITSCH. 1996. Functional as- for the Period June 1974–January 1975. 185 pp. sessment of five wetlands constructed to mitigate Rider College, Lawrenceville, NJ. wetland loss in Ohio, USA. Wetlands 16: 436–451. ZEDLER,J.B.AND J. C. CALLAWAY. 1999. Tracking WHIGHAM, D. F. 1999. Ecological issues related to wet- wetland restoration: do mitigation sites follow de- land preservation, restoration, creation and assess- sired trajectories? Restoration Ecol. 7: 69–73.

Appendix 1. Vascular plants of the Hamilton–Trenton–Bordentown Marsh. Taxa have been arranged in the following categories: vascular crytograms (lycopods, horsetails, and ferns), gym- nosperms, dicots, and monocots. Cultivated ephemerals are listed separately at the end. Alphabetical within each category are families and lower taxa. Each entry contains sci- entific name and authority. (Sources were Rhoads and Klein (1993) and, if a species is not recorded for Pennsylvania, then we used Gleason and Cronquist (1991). For Kyllinga we used Bryson, et al. (1997)). Additional notations may include: (1988) - species listed by Leck et al (1988); RM - reference marsh; CW - constructed wetland; sb - emerging from seed bank or spore bank soil samples; RM f are field species from Leck and Simpson 1987a, b; CW f are constructed wetland species recorded at the site (note that RM f and CW f designations do not preclude occurrences in other areas); and * indicates non-native species. Species that we have not seen ourselves are credited to their source, e.g., J. Arsenault, R. Blicharz, D. Snyder, or A. E. Schuyler (including specimens at the Academy of Natural Sciences of Philadelphia herbarium). We have also indicated, where appropri- ate, others who provided identifications. Species also listed by Bonaparte (1824) are in- dicated as [Bonaparte], by Abbott (1887), as [Abbott] and any additional species found in Abbott 1886, 1892, 1897, and 1898 as [Abbott ϩ], and by Shoemaker 1964 or 1965, as [Shoemaker]. (Other species found in Bonaparte (1824), Abbott (1886, 1887) or Shoe- maker (1964, 1965) appear in Appendix 2). Other comments considered to be of interest are also provided. For a list of New Jersey rare or endangered species see Table 3.

Ferns and Fern Allies Aspleniaceae LYCOPODIOPHYTA Asplenium platyneuron (L.) BSP. (1988). Isoetaceae Isoetes riparia Engelm. ex A. Braun. CW f. Several Blechnaceae plants were observed once during fall 1994 in a Woodwardia areolata (L.) T.Moore. (1988). RM sb. shallow depression near the east navigation bea- Woodwardia virginica (L.) Smith. RM sb only. Young con. plant from soil sample identified by J. Montgom- ery. Lycopodiaceae Huperzia lucidula (Michx.) Trevisan. One small plant Dennstaedtiaceae located near Spring Lake trails bridge. Did not Dennstaedtia punctilobula (Michx.) T.Moore. (1988). persist; area flooded due to beaver activity. RM sb. [Shoemaker]. Lycopodium obscurum L. Extensive patches near Stur- Pteridium aquilinum (L.) Kuhn. Bordentown Bluffs geon Pond and at N. Community Park. along Crosswicks Creek. [Abbottϩ].

Selaginellaceae Dryopteridaceae Selaginella apoda (L.) Fern. Oberved once on a path Athyrium filix-femina (L.) Roth. (1988). RM sb. [Shoe- in woods near Sturgeon Pond east. maker]. EQUISETOPHYTA Cystopteris tenuis (Michx.) Desv. Between grouted Equisetaceae rocks of old bridge abutment near Yardville. Equisetum arvense L. (1988). CW f. [Abbott] [Shoe- Dryopteris carthusiana (Villars) H.P. Fuchs. (1988). maker]. Dryopteris intermedia (Muhl. ex Willd.) A.Gray Equisetum fluviatile L. (1988). CW f. Infrequent. Dryopteris marginalis (L.) A.Gray Equisetum hyemale L. var. affine x laevigatum. Loca- Dryopteris sp. (1988). RM sb. tion on Duck Is. destroyed by highway construc- Matteuccia struthiopteris (L.) Todaro. (1988). Infre- tion. quent. Equisetum sp. (1988) RM sb. Onoclea sensibilis L. (1988). RM sb, CW sb, RM f, CW f. Most common fern in soil samples (Leck PTERIDOPHYTA and Simpson 1987b). [Shoemaker]. 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 341

Polystichum acrostichoides (Michx.) Schott. (1988). Acer saccharinum L. (1988). CW sb, RM f, CW f. [Abbottϩ]. Acer saccharum Marshall. Several sizable trees along Woodsia obtusa (Spreng.) Torr. Growing between road from Spring Lake to Watson Woods, likely rocks of abandoned bridge abutment on Delaware planted. River north of PSE&G. Amaranthaceae Ophioglossaceae Amaranthus cannabinus (L.) Sauer (1988). RM sb, Botrychium dissectum Spreng. (1988). CW f. D & R CW sb, RM F, CW f. Canal tow path on Duck Is. and forested island *Amaranthus retroflexus L. CW f. near RM. *Froelichia gracilis (Hook.) Moq. (1988). CW f. Botrychium virginianum (L.) Swartz. (1988). Spring Lake trails. Anacardiaceae Rhus copallina L. (1988). Osmundaceae Rhus glabra L. Infrequent. (1988). Noted in 1987 Osmunda cinnamomea L. (1988). [Abbottϩ] [Shoe- along road near brick factory. [Shoemaker]. maker]. Rhus typhina L. (1988). CW f [Shoemaker]. Osmunda claytoniana L. (1988). Infrequent. [Shoe- Toxicodendron radicans (L.) Kuntze. (1988). CW sb, maker]. RM f, CW f. [Abbott] [Shoemaker]. Osmunda regalis L. (1988). Infrequent; several popu- lations at Yardville - Delaware & Raritan Canal Apiaceae State Park. Cicuta bulbifera L. (1988). CW sb, CW f. Cicuta maculata L. (1988). RM sb, CW sb, RM f, CW f. Thelypteridaceae Cryptotaenia canadensis (L.) DC. Thelypteris noveboracensis (L.) Nieuwl. (1988). CW *Daucus carota L. (1988). CW f. f. [Shoemaker]. Heracleum lanatum Michx. (1988). RM f. Roadside at Thelypteris palustris Schott (1988). RM sb. Watson Woods, along railroad, and Lamberton Seed Plants Road. *Hydrocotyle sibthorpioides Lam. Bordentown beach, PINOPHYTA (Gymnosperms) Trenton boat launch. Cupressaceae Osmorhiza claytonii (Michx.) C.B.Clarke Chamaecyparis thyoides (L.) BSP. Several trees at Osmorhiza longistylis (Torr.) DC. (1988). edge of marsh near Bordentown Water Authority *Pastinaca sativa L. building, east of bridge at Rt. 206 - Crosswicks Ptilimnium capillaceum (Michx.) Raf. (1988, Arsen- Creek. [Abbott refers to ‘cedars along river- ault). banks’]. Stumps visible at low tide along Watson Sanicula odorata (Raf.) Pryer & Phillippe Creek may be this species. Sium sauve Walt. (1988). RM sb, CW sb, RM f, CW f. Juniperus virginiana L. Local, near Spring Lake. [Bonaparte] [Abbottϩ]. Apocynaceae Apocynum androsaemifolium L. (1988). Pinaceae Apocynum cannabinum L. (1988). CW sb, CW f. *Picea abies (L.) Karst. South tip Duck Island. *Vinca minor L. At abandoned homesites. [Shoemak- Pinus rigida P.Mill. (1988). Infrequent, Bordentown er]. Bluffs. Pinus strobus L. CW f. Infrequent. [Shoemaker]. Aquifoliaceae Tsuga canadensis (L.) Carr. Infrequent, Bordentown Ilex opaca Ait. (1988). Infrequent. Bluffs; many trees killed by wooly adelgid infes- Ilex verticillata (L.) A.Gray. (1988). CW sb. [Abbott] tation. A notable tree was 45 cm DBH (2004). [Shoemaker]. Taxaceae Araliaceae *Taxus spp. (1988). Planted at abandoned homesites Aralia nudicaulis L. Bordentown Bluffs, Watson (near Spring Lake and south tip of Duck Is. near Woods. Delaware & Raritan Canal). *Hedera helix L. Extensive patches at Bordentown Bluffs and bluffs near Rowan St. storm drain. MAGNOLIOPHYTA (Flowering Plants) MAGNOLIOPSIDA (Dicotoleydons) Aristolochiaceae Acanthaceae Asarum canadense L. (1988). CM f. Extensive patch Justicia americana (L.) Vahl. (1988). CW f. Several on floodplain east side of Duck Creek. populations along Delaware River, vicinity of CW, Trenton boat launch, and I 295 scenic overlook; Asclepiadaceae upper intertidal on cobble. Asclepias incarnata L. (1988). CW f. [Shoemaker]. Many plants along edge of Spring Lake in 2004. Aceraceae Asclepias syriaca L. (1988). CW f. [Shoemaker]. Acer negundo L. (1988). CW sb, CW f. [Abbott] [Shoemaker]. Asteraceae *Acer palmatum Thunb. Escaped, woods near Yard- *Achillea millefolium L. (1988). CW f. [Abbottϩ]. ville storm drain. Ambrosia artemisiifolia L. (1988). CW sb, CW f. *Acer platanoides L. (1988). Ambrosia trifida L. (1988). RM sb, RM f, CW f. Large *Acer pseudoplatanus L. (1988). [Shoemaker]. populations can occur on tidal channel levees. Acer rubrum L. (1988). RM f, CW f. [Abbott] [Shoe- Antennaria plantaginifolia (L.) Richardson. Site on maker]. Duck Is. destroyed during excavation for con- 342 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

structed wetland; one site on Bordentown Bluffs (1992, 1993) where Sorgastrum nutans was pre- near Stanton Ave. in 2004. sent. *Anthemis arvensis L. Eupatorium aromaticum L. (2 Aug. 1992). *Anthemis cotula L. (1988). Eupatorium dubium Willd. ex Poir. (1988). RM sb, *Anthemis tinctoria L. CW f. RM f, CW f. *Arctium minus (Hill.) Bernh. Eupatorium fistulosum Barratt. CW f. *Artemisia annua L. CW f. Eupatorium hyssopifolium L. (1988). CW f. *Artemisia vulgaris L. (1988). CW sb, CW f. Eupatorium leucolepis (DC.) Torr &. A.Gray. Near Aster cordifolius L. CW f. Sturgeon Pond. Aster divaricatus L. sensu lato (1988). Eupatorium maculatum L. CW f. Aster dumosus L. sensu lato.CWf. Eupatorium perfoliatum L. (1988). CW sb, CW f. [Ab- Aster ericoides L. CW sb, CW f. bott]. Aster fragilis Willd. var. fragilis (including A. race- Eupatorium purpureum L. (1988). RM f, CW f. [Shoe- mosus and A. vimineus). (1988). CW sb, CW f. maker]. Aster lanceolatus Willd. ssp. simplex (Willd.) Eupatorium rotundifolium L. A.G.Jones (1988). CW f. Eupatorium rugosum Houtt. (1988). CW sb, CW f. Aster longifolius Lam. sensu lato (A. praealtus Poir. Eupatorium serotinum Michx. CW sb, CW f. Domi- identified by J. Meyer). nant on Elatine Is. upland several years including Aster pilosus Willd. (1988). RM sb, CW sb, RM f, CW f. 2003. Also, during 1995-1997 on CW north marsh Aster puniceus L. (1988). CW sb, CW f. near channel. Baccharis halimifolia L. (1988). RM sb, CW f. Euthamia graminifolia (L.) Nutt. (1988). RM sb, CW Bidens bidentoides (Nutt.) Britt.. (1988). CW sb, CW sb, CW f. f. May be locally numerous, upper intertidal along *Galinsoga quadriradiata Ruiz & Pavon. CW f. Delaware River where there is disturbance; Tren- Gnaphalium obtusifolium L. (1988). RM sb, RM f, ton boat launch on Lamberton Road and else- CW f. where. Gnaphalium uliginosum L. CW sb, CW f. Bidens bipinnata L. CW f. Helenium autumnale L. (1988). CW sb, CW f. [Ab- Bidens cernua L. CW f. bott]. Bidens comosa (A.Gray) Wieg. (1988, Schuyler). CW sb. Helianthus annuus L. (1988, Blicharz). Bidens connata MuhL. (1988). CW sb, CW f. Locally Helianthus decapetalus L. (1988). CW f. Helianthus tuberosus L. (1998). CW f. abundant, e.g., 2003 marsh near Sturgeon Pond Heterotheca subaxillaris (Lam.) Britt. & Rusby. CW f. east. *Hieracium caespitosum Dumort. (1988). CW f. Bidens coronata (L.) Britt. (1988) CW f. Hieracium gronovii L. Bidens discoidea (Torr. & Gray) Britt. *Hieracium piloselloides Vill. Bidens frondosa L. (1988). RM sb, CW sb, RM f, CW f. Hieracium venosum L. Hillside near railroad, Yard- Bidens laevis (L.) BSP. (1988). RM sb, CW sb, RM f, ville. CW f. *Hypochoeris radicata L. *Bidens polylepis S.F.Blake (1988). CW sb, CW f. Krigia biflora (Walt.) S.F.Blake. West trail, Spring Bidens tripartida L. CW sb. Lake. Cacalia atriplicifolia L.. Two plants found in 1992 and Krigia virginica (L.) Willd. (1988). CW f. [Abbott]. for several years, but not recently (2001, 2002), Lactuca biennis (Moench) Fern. Beneath railroad Crosswicks Creek floodplain southwest (up- bridge, Yardville; Watson Woods. stream) of railroad bridge at Yardville. Lactuca canadensis L. (1988). RM sb, RM f, CW f. *Carduus nutans L. Roadside on Duck Is. *Lactuca serriola L. CW f. *Centaurea maculosa Lam. (1988). CW f. Lactuca sp. CW sb. *Chrysanthemum leucanthemum L. (1988). *Matricaria perforata Merat. CW f. *Cichorium intybus L. CW f. Mikania scandens (L.) Willd. (1988). RM sb, CW sb, *Cirsium arvense (L.) Scop. (1988). CW f. RM f, CW f. Common; dominant at CW. [Abbott]. *Cirsium vulgare (Savi) Tenore. (1988). CW sb; CW Prenanthes trifoliolata (Cass.) Fern. Wooded hillside f. [Abbott]. D & R Canal State Park near railroad (Yardville). Conyza canadensis (L.) Cronq. (1988). RM sb, CW Rudbeckia hirta L. (1988). CW f. sb; CW f. Rudbeckia laciniata L. (1988). CW f. [Abbott]. *Coreopsis lanceolata L. Roadside near PSE&G on Senecio aureus L. (1988). [Abbott]. Lamberton Road. *Senecio vulgaris L. CW f. *Crepis tectorum L. Solidago caesia L. (1988). Wooded hillsides near Eclipta prostata (L.) L. CW sb, CW f. Yardville and near Rowan St. drain. Erechtites hieraciifolia (L.) Raf. ex DC. (1988). RM Solidago canadensis L. (1988). RM sb, CW sb, RM f, sb, CW sb, RM f, CW f. CW f. Erigeron annuus (L.) Pers. (1988). RM sb, CW sb, Solidago canadensis var. scabra L. Torr. & A.Gray. CW f. CW f. Erigeron philadelphicus L. CW f. Solidago gigantea Ait. CW f. Erigeron strigosus Muhl. ex Willd. Solidago juncea Ait. (1988). CW f. Eupatorium album L. var. subvenosum A.Gray. CW f. Solidago rugosa Ait. (1988). CW sb, CW f. [Shoemaker]. Solidago sempervirens L. (1988). CW f. Eupatorium altissimum L.. CW f. CW along road to *Sonchus asper (L.) Hill. CW f. beacon (1996) and near Sturgeon Pond west *Taraxacum laevigatum (Willd.) DC. 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 343

*Taraxacum officinale Weber (1988). CW sb, CW f. persisted for several years until Tide Pool Is. be- [ϭ Taraxacum dens-leonis, Abbottϩ]. came overgrown. *Tragopogon dubius Scop. Open area near beach at *Eriophila verna (L.) Chev. CW f. [Abbottϩ]. Stanton Ave. *Hesperis matronalis L. CW f. Verbesina alternifolia (L.) Britt. CW sb, CW f. Lepidium virginicum L. CW f. Vernonia noveboracensis (L.) Michx. CW sb, CW f. Lepidulum sp. (1988). Common along Delaware River upper intertidal *Nasturtium officinale R.Br. (1988). RM sb, RM f. shore and frequent on high marsh of CW. [Ab- Rorippa palustris (L.) Bess. CW sb, CW f. bottϩ]. *Sisymbrium altissimum L. Disturbed upland at little Xanthium strumarium L. (1988). CW f. mitigation site. *Sisymbrium officinale (L.) Scop. Disturbed upland at Balsaminaceae little mitigation site. Impatiens capensis Meerb. (1988). RM sb, CW sb, RM *Thlaspi arvense L. CW f. f, CW f. [Abbott]. Impatiens pallida Nutt. Duck Is. along Lamberton Buxaceae Road, north of PSE&G. *Pachysandra terminalis Siebold & Zucc. Berberidaceae Cabombaceae *Berberis thunbergii DC. (1988). Cabomba caroliniana A.Gray. (1988). [Shoemaker]. Podophyllum peltatum L. (1988). [Bonaparte] [Ab- Caesalpiniaceae bott]. Chamaecrista fasciculata (Michx.) Greene (1988). Betulaceae Chamaecrista nictitans (L.) Moench. (1988). CW f. *Alnus glutinosa (L.) Gaertn. CW sb, CW f. Gleditsia triacanthos L. CW f. [Bonaparte]. Alnus incana (L.) Moench. [Shoemaker]. Senna hebecarpa (Fern.) Irwin & Barneby. CW f; one Alnus serrulata (Dryand. ex Ait.) Willd. (1988). RM plant several years on D.O.T. Is. and also Elatine sb, RM f [Abbott]. Is. in 2004. Betula lenta L. Bordentown Bluffs. Callitrichaceae Betula nigra L. (1988). RM sb, CW sb, RM f, CW f. Callitriche heterophylla Pursh emend. Darby. (1988). [Abbott]. RM sb, RM f, CW f. Betula populifolia Marshall. (1988). RM sb, RM f. *Callitriche stagnalis Scop. CW f. Carpinus caroliniana Walt. (1988). Corylus americana Walt. [Abbott]. Campanulaceae Campanula aparinoides Pursh. Infrequent; alder swamp Bignoniaceae at Watson Woods and Yardville–D & R Canal Campsis radicans (L.) Seem. ex Bureau. [Abbott]. State Park. Catalpa bignonioides Walt. (1988). CW f. [Abbott]. Lobelia cardinalis L. (1988). CW sb, CW f. Regular *Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex along Duck Creek. [Abbott]. Steud. (1988). RM sb, CW sb, RM f, CW f. Lobelia inflata L. (1988). CW f. Boraginaceae Lobelia siphilitica L. (1988). CW sb, CW f. Infre- *Echium vulgare L. quent. Hackelia virginiana (L.) I.M.Johnston Triodanis perfoliata (L.) Nieuwl. (1988). CW f. Mertensia virginica (L.) Pers. ex Link. Crosswicks Cannabaceae Creek floodplain near Yardville. *Humulus japonicus Sieb. & Zucc. (1988). CW f. *Myosotis arvensis (L.) Hill. CW f. Myosotis laxa Lehm. (1988). RM sb, CW sb, RM f, Caprifoliaceae CW f. *Lonicera japonica Thunb. (1988). RM f, CW f. *Lonicera morrowii A.Gray. (1988). CW f. Brassicaceae Lonicera sempervirens L. Two locations at Borden- *Alliaria petiolata (Bieb.) Cavara & Grande. (1988). town Bluffs (along trail and ravine west of Stanton CW f. Ave. beach). [Abbottϩ]. *Arabidopsis thaliana (L.) Heynh. CW f. *Lonicera xylosteum L. (1988). Arabis laevigata (Muhl. ex Willd.) Poir. (1988). CW f. Sambucus canadensis L. (1988). RM f, CW f. [Abbott] Barbarea verna (P.Mill.) Aschers. [Shoemaker]. *Barbarea vulgaris R.Br. CW f. Viburnum acerifolium L. Bordentown Bluffs and N, *Brassica nigra (L.) W.D.J.Koch. CW f. Community Park. [Shoemaker]. *Camelina microcarpa Andrz. ex DC. Disturbed up- Viburnum dentatum L. (1988). RM sb, RM f, CW f. land at little mitigation site. [Abbott] [Shoemaker]. *Capsella bursa-pastoris (L.) Medic. (1988). [Bona- Viburnum prunifolium L. (1988). RM sb, RM f, CW parte]. f. [Abbott]. Cardamine bulbosa (Schreb. ex Muhl.) BSP. (1988). RM sb, RM f. Caryophyllaceae Cardamine concatenata (Michx.) O.E.Schulz. [Shoe- *Arenaria serpyllifolia L. CW f. maker]. *Cerastium semidecandrum L. CW f. Cardamine hirsuta L. CW f. *Dianthus armeria L. (1988). CW f. Cardamine pensylvanica Muhl. ex Willd. (1988). RM Paronychia fastigiata (Raf.) Fern. CW f. sb, CW sb, CW f. *Sagina japonica (Swartz) Ohwi. CW f. *Diplotaxis tenuifolia (L.) DC. CW f. One plant that Sagina procumbens L. CW f. 344 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Sagina sp. CW sb. Droseraceae *Saponaria officinalis L. (1988). CW f. [Abbottϩ]. Drosera rotundifolia L. (1988). RM sb only. *Scleranthus annuus L. CW f. Ebenaceae *Silene alba (Mill.) Krause Diospyros virginiana L. (1988). CW f. Crosswicks Silene antirrhina L. [Bonaparte]. Creek floodplain, N. Community Park. [Abbott]. Silene caroliniana Walt. Small population near rail- road northeast of railroad bridge at Yardville. Elaeagnaceae *Silene noctiflora L. *Elaeagnus umbellata Thunb. CW f. Silene stellata (L.) Ait.f. CW f. Hillsides near Yard- ville, Spring Lake, and CW on Tide Pool Is. Elatinaceae *Spergula arvensis L. CW f. Elatine americana (Pursh) Arn. CW f, in ‘tidal’ pools; Stellaria longifolia Muhl. ex Willd. (1988). observed for several years but not in 2004. *Stellaria media (L.) Vill. CW f. [Abbottϩ]. Ericaceae Epigaea repens L. Infrequent, Bordentown Bluffs and Celastraceae ϩ *Celastrus orbiculatus Thunb. (1988). CW f. along trail at N. Community Park. [Abbott ]. *Euonymus alatus (Thunb.) Sieb. (1988). Kalmia latifolia L. Forming thickets along Borden- town Bluffs, also N. Community Park. [Abbott]. Ceratophyllaceae Rhododendron (?) canescens (Michx.) Sweet. South Ceratophyllum demersum L. (1988). CW f. end of Duck Is. near Bordentown. Rhododendron maximum L. Forming thickets along Chenopodiaceae Bordentown Bluffs. [Bonaparte]. Atriplex prostrata Bouder ex DC. CW sb, CW f. Rhododendron periclymenoides (Michx.) Shinners. *Chenopodium album L. (1988). CW sb, CW f. Ocassional plants along Crosswicks Creek, com- *Chenopodium ambrosioides L. (1988). CW sb, CW f. mon at N. Community Park. [Abbottϩ]. *Chenopodium pumilio R.Br. Vaccinium corymbosum L. (1988). *Cycloloma atriplicifolium (Spreng.) Coult. (1988). Vaccinium pallidum Ait. Bordentown Bluffs area. Cistaceae Vaccinium stamineum L. Bordentown Bluffs. [Ab- Lechea villosa Ell. (1988). bottϩ]. Clethraceae Euphorbiaceae Clethra alnifolia L. (1988). [Abbott]. Acalypha rhomboidea Raf. CW sb, CW f. Acalypha virginica L. (1988). CW sb, CW f. Clusiaceae Chamaesyce maculata (L.) Small. CW sb, CW f. Hypericum gentianoides (L.) BSP Chamaesyce nutans (Lag.) Small Hypericum mutilum L. (1988). CW sb, CW f. Euphorbia corollata L. (1988). Open upland along *Hypericum perforatum L. (1988). CW f. Spring Lake trails; area now has tree canopy. Hypericum punctatum Lam. CW sb, CW f Poinsettia dentata (Michx.) Klotzsch & Garcke. Along Triadenum virginicum (L.) Raf. (1988). RM sb, CW sb. railroad at Yardville. Convolvulaceae Fabaceae Calystegia sepium (L.) R.Br. sensu lato. (1988). CW Amorpha fruticosa L. (1988). CW sb, CW f. Common f. [Abbott]. colonizer along wetland upland edges of CW. *Convolvulus arvensis L. CW f. Amphicarpaea bracteata (L.) Fern. (1988). CW f. [Ab- *Ipomoea hederacea Jacq. bott]. Ipomoea pandurata (L.) G.F.W. Mey. Apios americana Medic. (1988). CW f. Locally com- Cornaceae mon. [Abbott]. Cornus alternifolia L.f. Northern Community Park. *Coronilla varia L. (1988). CW f. Cornus amomum P.Mill. (1988). RM sb, RM f, CW f. Crotalaria sagittalis L. Cornus florida L. (1988). CW f. [Bonaparte] [Abbott]. Desmodium canadense (L.) DC. Cornus racemosa Lam. Desmodium canescens (L.) DC. (1988, Arsenault). Cornus sericea L. Desmodium glabellum (Michx.) DC. Cornus sp. CW sb. Desmodium paniculatum (L.) DC. Identified by W. Standaert. Crassulaceae *Kummerowia stipulacea (Maxim.) Schindler *Sedum acre L. *Kummerowia striata (Thunb.) Shindl. CW f. *Sedum sarmentosum Bunge. CW f. *Lathyrus latifolius L. *Sedum purpureum (L.) J.A. Schultes (or S. telephium Lespedeza capitata Michx. (1988). CW f. L.) *Lespedeza cuneata (Dum.-Cours.) G.Don. CW f. Lespedeza intermedia Cucurbitaceae (S.Wats.) Britt. (1988). CW f. Lespedeza violacea (L.) Pers. Sicyos angulatus L. CW f. *Lotus corniculatus L. CW f. Cuscutaceae *Medicago lupulina L. Cuscuta compacta Juss. ex Choisy. Sturgeon Pond, *Medicago sativa L. identified by D. Snyder (Aug. 1993). *Melilotus alba Medic. (1988). CW f. Cuscuta gronovii Willd. ex Schultz (1988). RM sb, *Melilotus officinalis (L.) Pallas. (1988). CW f. CW sb, RM f, CW f. [Abbott]. Robinia pseudoacacia L. (1988). CW sb, CW f. Com- Cuscuta polygonorum Engelm. CW f. Infrequent; no mon colonizer edges of CW and islands. [Bona- information about persistence. parte] [Abbott]. 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 345

Strophostyles helvola (L.) Ell. CW sb, CW f. Aesculus hippocastanum L. Duck Is., infrequent. *Trifolium arvense L. (1988). CW f. *Trifolium aureum Pollich (1988). CW f. Hydrangeaceae *Trifolium campestre Schreb. Hydrangea arborescens L. Steep wooded hillside near *Trifolium dubium Sibth. CW f. Blacks Creek, west of Bordentown. *Trifolium hybridum L. CW f. *Philadelphus coronarius L. (1988). Bordentown *Trifolium pratense L. (1988). CW f. [Abbott]. Bluffs. *Trifolium repens L. (1988). CW sb, CW f. [Bonapar- te]. Hydrophyllaceae *Vicia sativa L. CW f. Hydrophyllum virginianum L. Floodplain along Wat- *Vicia tetrasperma (L.) Schreb. son Creek. *Wisteria floribunda (Willd.) DC. (1988). Juglandaceae Fagaceae Carya cordiformis (Wang.) K.Koch (1988). Castanea dentata (Marshall) Borkh. Several plants, Carya glabra (P.Mill.) Sweet [Abbott]. Bordentown Bluffs and N. Community Park. [Ab- Carya ovata (P.Mill.) K.Koch. RM f. [Abbott]. bott]. Carya tomentosa (Lam. ex Poir.) Nutt. [Bonaparte]. Fagus grandifolia Ehrh. (1988). CW f. [Abbott] [Shoe- Juglans nigra L. CW f. [Bonaparte] [Abbott]. maker]. Quercus alba L. [Bonaparte] [Abbott] [Shoemaker]. Lamiaceae Quercus bicolor Willd. [Bonaparte] [Abbott]. Agastache nepetoides (L.) Kuntze. Old road near Quercus marilandica Muenchh. Identified by J. Meyer. sludge lagoon at Hamilton water treatment plant. Quercus michauxii Nutt. [ϭ Lophanthus nepethoides Benth. Abbottϩ]. Quercus montana Willd. [Bonaparte]. Agastache scrophulariifolia (Willd.) Kuntze (1988) Quercus muhlenbergii Engelm. Observed along east end of old road by sludge Quercus palustris Muenchh. (1988). RM f, CW f. lagoon at Hamilton Water Treatment Plant; hill- [Bonaparte] [Abbottϩ] [Shoemaker]. side near Sturgeon Pond east. Quercus phellos L. (1988). Sunny Pond near Spring Collinsonia canadensis L. Bordentown Bluffs. Lake spillway; also one tree along road from Dracocephalum parviflorum Nutt. CW f. Duck Island to Spring Lake in 1986. [Abbottϩ *Elsholtzia ciliata (Thunb.) Hylander. CW f; noted noted a plantation of this species]. near northernmost inlet in 1996; frequent along Quercus rubra L. (1988). RM f. [Bonaparte] [Shoe- edges of several islands in 2003. maker]. *Glechoma hederacea L. (1988). CW f. Quercus stellata Wang. [Abbottϩ]. Hedeoma pulegioides (L.) Pers. CW sb. Quercus velutina Lam. *Lamium amplexicaule L. (1988, Blicharz). CW f. * Lamium purpureum L. Fumariaceae *Leonurus cardiaca L. (1988). CW f. Roadsides at Corydalis flavula (Raf.) DC. (1988). CW f. [Abbottϩ]. Watson Woods. Dicentra cucullaria (L.) Bernh. (1988). Abbott (1886) Lycopus americanus Muhl. ex Bart. (1988). CW sb, called this ‘Eardrops’, perhaps more politically CW f. correct at the time than ‘Dutchman’s Breeches’. *Lycopus europaeus L. CW sb, CW f. [Abbottϩ] [Shoemaker]. Lycopus rubellus Moench. CW sb, CW f. Gentianaceae Lycopus virginicus L. (1988). RM sb, CW sb, RM f, Bartonia paniculata (Michx.) Muhl. Near Sturgeon CW f. Mentha arvensis Pond east. L. CW sb, CW f. Occasional in tidal Bartonia virginica (L.) BSP. N. Community Park. areas. Monarda fistulosa L. Geraniaceae *Nepeta cataria L. Erodium cicutarium (L.) L’Her. ex Soland. *Perilla frutescens (L.) Britt. CW f; at north inlet in Geranium carolinianum L. CW f. [Bonaparte]. 1996 and on several islands in 2003. Geranium maculatum L. Bordentown Bluffs and Cros- *Prunella vulgaris L. (1988). CW sb, CW f. swicks Creek floodplain at Yardville. [Abbottϩ]. Pycnanthemum tenuifolium Schrad. (1988). Pycnanthemum verticillatum (Michx.) Pers. Haloragaceae Pycnanthemum virginianum (L.) Durand & Jackson ex Myriophyllum heterophyllum Michx. (1988, Schuyler). B.L.Robins & Fern. (1988, Blicharz). *Myriophyllum spicatum L. (1988). CW sb, CW f. Salvia lyrata L. [Abbott]. Scutellaria galericulata L. (1988). CW sb, CW f. Proserpinaca palustris L. Sturgeon Pond and contig- Scutellaria integrifolia L. (1988). CW f. uous marsh, near landfill. Scutellaria lateriflora L. (1988). RM sb, CW sb, RM Hamamelidaceae f, CW f. Hamamelis virginiana L. [Abbottϩ]. Stachys palustris L. (1988, Schuyler). CW f. Liquidambar styraciflua L. (1988). RM sb, RM f, CW Stachys tenuifolia Willd. var hispida (1988). ?S3.RM f. [Bonaparte] [Abbott]. f, CW f. Teucrium canadense L. (1988). CW f. Hippocastanaceae Trichostema dichotomum L. (1988). CW f. Aesculus flava Ait. Wooded hillside at Point Breeze overlooking Crosswicks Creek; regeneration is ap- Lauraceae parent. Lindera benzoin (L.) Blume (1988). CW f. [Abbottϩ]. 346 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Sassafras albidum (Nutt.) Nees. (1988). RM f, CW f. Comptonia peregrina (L.) Coult. [Bonaparte] [Abbottϩ]. Nelumbonaceae Lentibulariaceae Nelumbo lutea (Willd.) Pers. Schuyler, herbarium only. Utricularia gibba L. Sturgeon Pond and Spring Lake. [Abbott]. A photograph (Abbott 1892) shows this Utricularia intermedia Hayne. (1988, Schuyler). growing in a garden tub. Utricularia macrorhiza Leconte (U. vulgaris L.). (1988, Schuyler). Nyctaginaceae Utricularia radiata Small. At west end of Spring Lake, Mirabilis nyctaginea (Michx.) MacM. Duck Is., along flowering late April 2002. Lamberton Road. Limnanthaceae Nymphaeceae Floerkea prosperpinacoides Willd. Common, Cros- Nuphar lutea (L.) Sibth. & Smith sensu lato. (1988). swicks Creek floodplain near Yardville. RM sb, CW sb, RM f, CW f. [Abbott] [Shoemak- er]. Loganiaceae Nymphaea odorata Ait. (1988). Once common (Clyde *Buddleja davidii Franch. Common along railroad on Quin, pers. comm.); only observed in Sturgeon Duck Is. Pond west. Plantings in CW (Tide Pool Is.) ap- Lythraceae parently extirpated by 2004. [Abbottϩ] [Shoe- Decodon verticillatus (L.) Ell. Beaver pond west of the maker]. Spring Lake spillway, noted in 2002. Nyssaceae th Lythrum alatum Pursh (1988). RM f. Ranked 11 (of Nyssa sylvatica Marshall. (1988). CW sb, RM f, CW 22) in importance in open marsh vegetation by f. [Abbott] [Shoemaker]. Whigham and Simpson (1975), but has not been observed by authors. Oleaceae *Lythrum salicaria L. (1988). RM sb, CW sb, RM f, Fraxinus americana L. (1988). RM sb, RM f, CW f. CW f. Not recorded by Whigham and Simpson [Abbottϩ]. (1975) in open marsh or shrub forest plots. Gen- Fraxinus pennsylvanica Marshall (1988). CW sb, RM erally infrequent except in CW. f, CW f. *Ligustrum obtusifolium Sieb. & Zucc. Magnoliaceae *Ligustrum vulgare L. (1988). CW f. [Shoemaker]. Liriodendron tulipifera L. (1988). RM sb, CW sb, RM f, CW f. [Bonaparte] [Abbott]. Onagraceae Magnolia tripetala L. Along Bordentown Bluffs and Circaea lutetiana L. ssp. canadensis (L.) Ascher & at Divine Word Missionaries easement in woods Magnus (1988). near Park Street. Epilobium coloratum Biehler (1988). RM sb, CW sb, Magnolia virginiana L. One sizable tree at Abbott’s RM f, CW f. swamp, and a few small trees at N. Community Ludwigia alternifolia L. (1988). CW sb, CW f. Park. [Bonaparte] [Abbott]. Ludwigia palustris (L.) Ell. (1988). RM sb, CW sb, Malvaceae RM f, CW f. *Abutilon theophrastii Medic. CW f. Ludwigia peploides (Kunth) Raven. (1988). CW f. Ob- Hibiscus moscheutos L. (1988). RM sb, CW sb, RM served on Elatine Is. in CW during fall 2002; did f, CW f. [Abbottϩ] [Shoemaker]. not persist. Common at Sturgeon Pond. *Hibiscus trionum L. CW f. Oenothera biennis L. (1988). RM sb, CW sb, CW f. *Malva neglecta Wallr. CW f. [Abbott]. Oenothera perennis L. Bordentown Bluffs, sandy area Melastomataceae near beach at Stanton Ave. Rhexia virginica L. (1988). Infrequent. Orobanchaceae Menispermaceae Epifagus virginiana (L.) Bart. Menispermum canadense L. CW f. Orobanche uniflora L. Uncommon; found once on Mimosaceae abandoned road to forested island near Hamilton *Albizia julibrissin Durazz. CW f. water treatment plant and RM. Molluginaceae Oxalidaceae *Mollugo verticillata L. (1988). CW sb, CW f. Oxalis stricta L. (1988). CW f. Oxalis violacea L. Bordentown Bluffs; observed in Monotropaceae 1992, but not in 2004. [Bonaparte]. Monotropa uniflora L. (1988). Infrequent Sturgeon Pond area; Forest Is. near RM, Watson Woods. Papaveraceae *Chelidonium majus L. (1988). Moraceae *Broussonetia papyrifera (L.) Vent. (1988). Abbott’s Phytolaccaceae homesite area. Phytolacca americana L. (1988). CW sb, CW f. [Ab- *Maclura pomifera (Raf. Ex Sarg.) Schneid. One small bott]. plant near Trenton boat launch. Plantaginaceae *Morus alba L. (1988). CW f. Plantago aristata Michx. (1988). CW f. Morus rubra L. CW f. [Bonaparte]. *Plantago lanceolata L. (1988). CW f. [Bonaparte]. Myricaceae *Plantago major L. CW sb. 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 347

Plantago rugelii Decne. (1988). CW f. *Ranunculus ficaria L. CW f. Well established along Delaware River in 2003, especially north of little Platanaceae mitigaton site; first occurrence not noted. First ob- Platanus occidentalis L. (1988). CW sb, CW f. [Bon- served in CW in 2001. Not recorded along Cros- aparte] [Abbott] [Shoemaker]. swicks Creek. Polemoniaceae Ranunculus hispidus Michx. CW f. Phlox paniculata L. Ranunculus longirostris Godr. (1988, Schuyler, her- barium only). Polygonaceae *Ranunculus sceleratus L. (1988). RM sb, RM f, CW f. Polygonella articulata (L.) Meisn. CW f. Thalictrum pubescens Pursh (1988). RM sb, RN f, CW Polygonum amphibium L. (1988). RM f, CW f. Stur- f. [Abbottϩ] [Shoemaker]. geon Pond west, Spring Lake; CW on Abbott Is. in 2004. Rosaceae Polygonum arifolium L. (1988). RM sb, CW sb, RM Agrimonia parviflora Ait. CW f. f, CW f. Amelanchier arborea (Michx.f.) Fern. Bordentown *Polygonum aviculare L. CW f. Bluffs. *Polygonum caespitosum Blume. (1988). CW sb, CW f. Aronia melanocarpa (Michx.) Ell. Bordentown Bluffs Poygonum careyi Olney. CW sb. along Crosswicks Creek. Polygonum cilinode Michx. CW f. Crataegus crus-galli L. sensu lato [Abbottϩ]. *Polygonum cuspidatum Sieb. & Zucc. (1988). CW f. Crataegus intricata Lange sensu lato. Identified by J. *Polygonum hydropiper L. CW sb, CW f. Meyer. Polygonum hydropiperoides Michx. (1988). CW sb, *Duchesnea indica (Andr.) Focke CW f. Fragaria virginiana P.Mill. Polygonum lapathifolium L. (1988). CW sb, CW f. Geum canadense Jacq. (1988). CW f. *Polygonum orientale L. (1988, Schuyler). Geum laciniatum Murr. (1988, Arsenault). CW f. Polygonum pensylvanicum L. (1988). CW sb, CW f. *Malus pumila P.Mill. Infrequent, Crosswicks Creek *Polygomum perfoliatum L. CW f, a few plants at floodplain. [Shoemaker]. north inlet 2001, 2002, but not 2003). Physocarpus opulifolius (L.) Maxim. (1988). RM sb, *Polygonum persicaria L. CW sb, CW f. RM f, CW f. Poygonum punctatum ElL. (1988). RM sb, CW sb, Potentilla norvegica L. (1988). RM sb, CW sb, CW f. RM f, CW f. *Potentilla recta L. Polygonum sagittatum L. (1988). RM sb, CW sb, RM Potentilla simplex Michx. (1988). CW f. f, CW f. Prunus serotina Ehrh. (1988). RM f, CW f. [Bonapar- Polygonum scandens L. (1988). CW f. te] [Abbottϩ]. Polygonum setaceum Baldw. ex Ell. S2. CW sb, CW f. Prunus serulata (?) One small tree at the edge of Ab- Polygonum virginianum L. (1988). RM f, CW f. bott’s swamp. *Rumex acetosella L. CW f. *Pyrus communis L. Infrequent, Crosswicks Creek Rumex altissimus A.Wood. CW sb, CW f. floodplain. *Rumex crispus L. (1988). CW f. *Rosa multiflora Thunb. ex Murr. (1988). CW f. *Rumex obtusifolius L. (1988). CW sb, CW f. Rosa palustris Marshall (1988). RM f. Rumex triangulivalvis (Danser) Rech.f. *Rosa spinosissima. One bush, south tip of Duck Is. Rubus allegheniensis Porter (ϭRubus pensilvanicus Portulacaceae Poir.) (1988). CW f. [Bonaparte] [Abbott]. Claytonia virginica L. (1988). CW f. [Abbott] [Shoe- Rubus flagellaris Willd. sensu lato [Shoemaker]. maker]. Rubus hispidus L. sensu lato. Portulaca oleracea L. CW f. *Rubus laciniatus Willd. Primulaceae Rubus occidentalis L. [Shoemaker]. *Anagallis arvensis L. Rubus odoratus L. CW f; did not persist. Lysimachia ciliata L. (1988). RM sb, CW sb, RM f, *Rubus phoenicolasius Maxim. (1988). CW f. CW f. Spiraea latifolia (Ait.) Borkh. (1988). CW sb. *Lysimachia nummularia L. CW f. Spiraea tomentosa L. (1988). Lysimachia quadrifolia L. Rubiaceae Lysimachia terrestris (L.) BSP. CW f. Cephalanthus occidentalis L. (1988). RM sb, CW sb, Pyrolaceae CW f. [Abbott]. Chimaphila maculata (L.) Pursh. (1988). Infrequent. Diodia teres Walt. CW f. Galium aparine L. (1988). CW f. [Bonaparte] Ranunculaceae Galium obtusum Bigel. (1988). RM f, CW f. ϩ Anemone quinquefolia L. [Bonaparte] [Abbott ] Galium palustre L. Identified by W. Standaert and P. [Shoemaker]. Cooney. Caltha palustris L. (1988). Locally common in tidal Galium tinctorium L. (1988). CW sb. areas near railroad (w) and Bordentown water Galium triflorum Michx. plant. [Abbott]. Hedyotis caerulea (L.) Hook. (1988). CW f. [Bona- *Clematis terniflora DC. (1988). CW sb, CW f. Com- parte] [Abbott] [Shoemaker]. mon along upland edges near Delaware River. Mitchella repens L. Ranunculus abortivus L. (1988). CW f. *Ranunculus bulbosus L. Lawn near edge of bluff at Rutaceae Point Breeze. [Abbottϩ]. *Phellodendron japonicum Maxim. One tree, Ͼ 10 cm 348 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

DBH along trail near Sturgeon Pond east Simaroubaceae (chopped down). *Ailanthus altissima (P.Mill.) Swingle (1988). CW sb, Ptelea trifoliata L. CW f. Several locations on Duck CW f. [Shoemaker]. Is.: open area north of CW, road to navigation beacon, and upland edge of west marsh, Lamber- Solanaceae ton Road. *Datura stramonium L. CW f. [Abbott]. *Lycium barbarum L. Near Sturgeon Pond during Salicaceae highway construction. Identified by D. Snyder. Populus balsaminifera L. Physalis heterophylla Nees. CW f. Populus deltoides Bartr. ex Marsh. Physalis subglabrata Mackenzie & Bush Populus grandidentata Michx. (1988). CW f. Solanum carolinense L. CW f. Populus heterophylla L. (1988). RM sb. One tree lo- *Solanum dulcamara L. (1988). [Abbott]. cated from canoe along tidal channel near brick Solanum nigrum L. (1988). CW sb, CW f. factory. Staphyleaceae *Populus nigra L. Staphylea trifolia L. Common, Crosswicks Creek Populus tremuloides Michx. (1988). floodplain. *Salix babylonica L. Planted at Spring Lake. [Abbott]. Salix bebbiana Sarg. (1988). RM sb. Tiliaceae Salix discolor Muhl. Tilia americana L. (1988). CW f. [Abbott]. Salix eriocephala Michx. (1988). RM sb, RM f. Salix exigua Nutt. CW f. Ulmaceae Celtis occidentalis L. (1988). *Salix fragilis L. (1988, Arsenault). ϩ Salix humilis Marshall (S. tristis?) (1988). Ulmus americana L. CW f. [Abbott ]. Salix hybrid (1988). RM sb. *Ulmus glabra Hudson. Identified by J. Kuser. Salix nigra Marshall. CW sb, CW f. [Shoemaker]. Ulmus rubra Muhl. (1988). RM f, CW f. Salix sericea Marshall. CW f. Urticaceae Boehmeria cylindrica (L.) Swartz (1988). RM sb, CW Sapindaceae sb, RM f, CW f. *Koelreuteria paniculata Laxm. Infrequent; along Laportea canadensis (L.) Weddell (1988). CW f. Delaware River. Pilea pumila (L.) A.Gray (1988). RM sb, CW sb, RM Saururaceae f, CW f. Saururus cernuus L. CW sb. Abbott’s swamp and Urtica dioica L. ssp. gracilis (Ait.) Seland. CW f; on along Crosswicks Creek upstream of railroad Tide Pool Is. bridge at Bordentown. Valerianceae Saxifragaceae *Valerianella locusta (L.) Betcke. CW f. [Bonaparte] Heuchera americana L. Bordentown Bluffs. [Abbott]. Penthorum sedoides L. (1988). CW sb, CW f. Verbenaceae Scrophulariaceae Verbena hastata L. CW sb, CW f. [Abbott]. Agalinus purpurea (L.) Pennell. [Abbottϩ]. Verbena urticifolia L. (1988). CW f. Chelone glabra L. (1988). RM sb, RM f, CW f. at Violaceae CW; did not persist. *Viola arvensis Murr. Gratiola neglecta Torr. (1988). Rm sb, CW sb, RN f, Viola lanceolata L. CW f. Viola macloskeyi Lloyd ssp. pallens (Banks ex DC) Linaria canadensis (L.) Dum.-Cours. (1988). CW f. M.S.Baker *Linaria vulgaris Hill. CW f. Viola eriocarpa Schwein. Lindernia dubia (L.) Pennell (1988). CW sb, CW f. Viola rafinesquii Greene (1988). CW f. [Shoemaker]. *Mazus pumilus (Burm.f.) Steenis. CW f. Edge Dela- Viola sororia Willd. (1988). RM sb, RM f. ware River near Trenton boat launch. Viola striata Ait. Edge of lawn at bluff, Point Breeze. Mimulus alatus Ait. (1988, Schuyler). CW f. Frequent at CW, 1995-1997, but did not persist. Vitaceae Mimulus ringens L. (1988). CW sb, CW f. Frequent *Ampelopsis brevipedunculata (Maxim.) Trautv. CW at CW 1995-1997, infrequent later; also white f. Identified by P. Cooney. form in 1995. Parthenocissus quinquefolia (L.) Planch. (1988). RM Scrophularia marilandica L. (1988). CW f. sb, RM f, CW f. [Abbott]. *Verbascum blattaria L. (1988). CW f. Vitis aestivalis Michx. (1988). RM f. *Verbascum lychnitis L. CW f. Duck Is., upland north Vitis labrusca L. (1988). CW f. [Abbott]. of CW; road to beacon. Vitis riparia Michx. (1988). CW f. *Verbascum thapsus L. (1988). CW sb, CW f. Vitis vulpina L. CW f. [Abbott][Shoemaker]. Veronica americana (Raf.) Schwein. ex Benth. CW f. Vitis sp. CW sb. Veronica anagallis-aquatica L. CW f. LILIOPSIDA (Monocotoleydons) Veronica arvensis * L. Acoraceae *Veronica hederifolia L. Woods near Yardville. Acorus calamus L. (1988). RM f, CW f. Extensive *Veronica longifolia L. (1988, Blicharz). stands in some high marsh areas, e.g., N. Com- Veronica peregrina L. (1988). RM sb, CW sb, RM f, munity Park. Not observed to produce seeds. CW f. Veronica persica Poir. N. Community Park. Agavaceae 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 349

Yucca flaccida Haw. Spring Lake trail. Carex laxiflora Lam. (ϭ Carex laxiculmis Schwein.) CW sb. Alismataceae Carex lupulina Muhl. ex Willd. (1988). CW f. Alisma plantago-aquatica L. var. parviflorum (Pursh) Carex lurida Wahlenb. CW sb, CW f. Torr. (1988). RM sb, CW sb, RM f, CW f. [Ab- Carex molesta Mackenzie ex Bright. CW f. bottϩ]. Carex normalis Mackenzie. CW f. Sagittaria graminea Michx. CW f. Carex pellita Muhl. ex Willd. Sagittaria latifolia Willd. (1988). RM sb, CW sb, RM Carex pensylvanica Lam. RM f. f, CW f. [Abbottϩ] [Shoemaker]. Carex projecta Mackenzie (1988). CW sb. Sagittaria rigida Pursh (1988, Schuyler). Carex radiata (Wahlenb.) Small (1988). CW f. Sagittaria subulata (L.) Buch (1988). CW f. Frequent; Carex rosea Schkuhr. ex Willd. (1988, Arsenault). CW on islands and north and east marshes, and Carex scoparia Schkuhr ex Willd. (1988). RM sb, CW along Delaware River where it occurs as turf near sb, RM f, CW f. low tide elevation. Carex squarrosa L. (1988). CW f. Carex sterilis Willd. CW sb, CW f. Araceae Carex stipata Muhl. Ex Willd. (1988). RM sb, CW sb, Arisaema dracontium (L.) Schott. CW f. One mature CW f. plant, Delaware River floodplain beneath trees, Carex straminea Willd. in Schkuhr (1988). near CW inlet. Carex stricta Lam. (1988). CW sb, CW f. Arisaema triphyllum (L.) Schott. (1988). [Shoemaker]. Carex swanii (Fern.) Mackenzie. (1988). RM sb, CW f. Orontium aquaticum L. (1988). About a dozen plants, Carex tribuloides Wahlenb. CW sb, CW f. upper intertidal on Crosswicks Creek (north Carex trichocarpa Muhl. ex Willd. Identified by W. bank); upstream and within sight of the Borden- Olson. town railroad bridge. One of two rarest species Carex trisperma Dewey noted by Abbott (1887). [Abbott]. Carex vulpinoidea Michx. (1988). CW sb, CW f. Peltandra virginica (L.) Schott & Endl. (1988). RM ϩ Cyperus bipartitus Torr. CW sb, CW f. Pseudo-vivip- sb, RM f, CW f. [Abbott ] [Shoemaker]. arous plantlets on greenhouse seed-bank speci- Symplocarpus foetidus (L.) Salisb. ex Nutt. (1988). ϩ men. Common along bluffs in seepage areas. [Abbott ] Cyperus dentatus Torr. (1988). CW sb, CW f. [Shoemaker]. C¸ yperus diandrus Torr. CW sb, CW f. Commelinaceae Cyperus echinatus (L.) A.Wood. (1988). CW f. *Commelina communis L. (1988). [Abbott]. Cyperus erythrorhizos Muhl. (1988). RM sb, CW sb, Tradescantia ohioensis Raf. S2. Duck Is. along rail- CW f. road. Cyperus esculentus L. CW sb. Tradescantia virginiana L. Frequent along railroads on Cyperus flavescens L. CW sb, CW f. Duck Is. and near Yardville. Cyperus lancastriensis Porter ex A.Gray. CW f. Cyperus lupulinus (Sprengel) Marcks (1988). CW sb, Cyperaceae CW f. Bordentown Bluffs, open sandy area near Bulbostylis capillaris (L.) C.B.Clarke. CW f. Duck Is. Stanton Ave. along railroad. *Cyperus microiria Steud. CW sb, CW f. Carex abscondita Mackenzie. CW f. Cyperus odoratus L. CW sb, CW f. Carex amphibola Steud. CW f. Cyperus retrofractus (L.) Torr. Carex annectans (Bickn.) Bickn. CW sb. Cyperus squarrosus L. CW sb. (Only sb). Carex argyrantha Tuckerman Cyperus strigosus L. (1988). RM sb, CW sb, RM f, Carex blanda Dewey. CW f. CW f. Carex brevior (Dewey) Mackenzie ex Lunell. CW f. Dulichium arundinaceum (L.) Britt. (1988). RM sb, Carex canescens L. CW sb, CW f. RM f, CW f. Carex caroliniana Schwein. CW f. Eleocharis acicularis (L.) Roemer & Schultes (1988). Carex cephalophora Muhl. ex Willd. CW f. CW sb, CW f. Carex comosa Boott. CW f. Eleocharis engelmannii Steud. CW sb, CW f. Carex crinita Lam. (1988). CW f. Eleocharis erythropoda Steud. CW sb. Carex cristatella Britt. CW f. Eleocharis obtusa (Willd.) Schultes (1988). CW sb, Carex davisii Schwein. & Torr. CW f. CW f. Carex debilis Michx. CW f. Eleocharis palustris (L.) Roemer & Schultes. CW sb, Carex festucacea Schkuhr ex Willd. CW f. Carex frankii Kunth. CW sb, CW f. Various terrestrial Eleocharis tenuis (Willd.) Schultes locations including upland north edge of CW, on Fimbristylis autumnalis (L.) Roemer & Schultes. CW islands, and near Bordentown beach. sb, CW f. Carex granularis Muhl. ex Willd. CW f. *Kyllinga gracillima Miq. CW sb, CW f. Carex grayi Carey. (1988). CW f. Rhynchospora capitellata (Michx.) Vahl Carex gynandra Schwein. Schoenoplectus fluviatilis (Torr.) Strong (1988). RM f, Carex hystericina Muhl. ex Willd. CW f. CW f. Carex interior Bailey Schoenoplectus pungens (americanus) (Vahl) Palla Carex intumescens Rudge (1988). (1988). CW sb, CW f. Carex lacustris Willd. (1988, Schuyler). Schoenoplectus purshianus (Fern.) Strong. CW f. Carex laevivaginata (Kukenth.) Mackenzie Pseudo-viviparous plantlets noted in the field. Carex laxiculmis Schwein. CW sb. Schoenoplectus smithii (A.Gray) Sojak. CW sb, CW f. 350 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Frequent during early succession at CW; did not and along the north and west marshes at Spring persist. Lake. Four previous state records (D. Snyder pers Schoenoplectus tabernaemontani (K.C.Gmel.) Palla. com.). CW sb, CW f. Scirpus atrocinctus Fern. CW f. Identified by A.E. Liliaceae Schuyler; did not persist. Allium canadense L. Woods at N. Community Park Scirpus atrovirens Willd. CW f. Field vivpary noted. and near brick factory. Scirpus cyperinus (L.) Kunth. (1988). RM sb, CW sb, *Allium vineale L. (1988). CW f. RM f, CW f. *Asparagus officinalis L. (1988). Scirpus expansus Fern. CW f. *Convallaria majalis L. Scirpus georgianus Harper. CW f. Pseudo-viviparous Erythronium americanum Ker-Gawl. (1988). [Abbott] plantlets noted in the field. [Shoemaker]. Scirpus hattorianus Makino. CW f. *Hemerocallis fulva (L.) L. [Shoemaker]. Scirpus polyphyllus Vahl. (1988). CW f. Pseudo-vivip- *Hosta sp. (probably H. ventricosa (Salisb.) Stearn). arous plantlets noted in the field. (1988). Lilium canadense L. Frequent along Crosswicks Creek Dioscoriaceae Lilium superbum L. (1988). CW f. [Abbott noted *Dioscorea batatas Decne. (1988). Near railroad plants Ͼ 2 m tall]. bridge at Bordentown. Maianthemum canadense Desf. Locally abundant. Dioscorea villosa L. (1988). RM sb, RM f. Medeola virginiana L. N. Community Park. *Narcissis pseudonarcissis L. At abandoned homesites Eriocaulaceae or planted. Near Watson House some single and Eriocaulon parkeri B.L.Robins (1988, Schuyler, her- doubled flowered plants, dating to 1915, were barium only). planted by Issac R. Van Zant (L. van Sant, pers. ϩ Hydrocharitaceae com.). [Abbott ]. *Egeria densa Planch. Small pond at N. Community *Ornithogalum umbellatum L. CW f. One plant noted Park in 2001. at Bordentwon Bluffs in 1993. [Bonaparte]. Elodea nuttallii (Planch.) St. John (1988). CW f. Polygonatum biflorum (Walt.) Ell. (1988). CW f. [Bon- ϩ *Hydrilla verticillata (L. f.) Royle. CW f. First found aparte] [Abbott ]. in 2003; dense growth in Tide Pool. Identified by Polygonatum biflorum (Walt.) Ell. var. commutatum A.E. Schuyler. (Schultes f.) Morong [Shoemaker]. Vallisneria americana Michx. (1988). CW f. Extensive Smilacina racemosa (L.) Desf. (1988). CW f. [Bona- beds along Delaware River from Bordentown to parte] [Shoemaker]. CW. [Abbott]. Uvularia sessilifolia L. (1988). Iridaceae Najadaceae *Iris pseudoacorus L. (1988). RM sb, RM f, CW f. Najas flexilis (Willd.) Rostk. & Schmidt. CW f. [Abbott]. Iris versicolor L. (1988). CW f. [Abbott] [Shoemaker]. Orchidaceae Sisyrinchium angustifolium P.Mill. (1988). CW f. Cypripedium acaule Ait. Once common, now only 5 [Shoemaker]. isolated, widely scattered non-flowering plants (3 Bordentown Bluffs, 1 N. Community Park, and 1 Juncaceae near Sturgeon Pond east). [Abbottϩ]. Juncus acuminatus Michx. (1988). CW sb, CW f. *Epipactis helleborine (L.) Crantz. Infrequent, near Pseudo-viviparous plantlets in the field.frequent. brick factory (2001, 2002); also N. Community Juncus brevicaudatus (Englem.) Fern. CW f. Identified Park. by A.E. Schuyler. On Elatine Is. in 1996, did not Platanthera lacera (Michx.) G.Don. One plant along persist. Spring Lake trail, woods near Beaver Point Juncus bufonius L. (2002). Juncus dichotomus Ell. CW f. Spiranthes cernua (L.) L.C.Rich. Open area near Stur- Juncus effusus L. (1988). RM sb, CW sb, RM f, CW f. geon Pond east (1991). Juncus secundus Beauv. ex Poir. CW f. Juncus tenuis Willd. (1988). RM sb. CW sb, RM f, Poaceae CW f. *Agrostis capillaris L. CW sb. Juncus torreyi Cov. CW f. Identified by A.E. Schuyler *Agrostis gigantea Roth. CW sb, RM f. in 1996; did not persist. Agrostis hyemalis (Walt.) BSP. CW f. Luzula multiflora (Ehrh.) Lej. (1988). Agrostis perennans (Walt.) Tuckerman. CW sb. *Agrostis stolonifera L. (1988). RM sb, CW f. Lemnaceae *Alopecurus pratensis L. Lemna minor L. (1988). RM sb, CW sb, RM f, CW f. Andropogon gerardii Vitman (1988). CW f. Lemna valdiviana Philippi. With other Lemnaceae, be- Andropogon ternarius Michx. CW f. On D.O.T Is; did low surface during cool weather (e.g. Dec.). Six not persist. previous state records (D. Snyder, pers. commun.). Andropogon virginicus L. (1988). RM sb, CW sb. Spirodela polyrhiza (L.) Schleid. (1988). CW f. *Anthoxanthum odoratum L. Wolffia columbiana Karst. (1988). [Shoemaker]. Aristida dichotoma Michx. (1988). Wolffia papulifera C. Thompson Aristida oligantha Michx. Wolffiella gladiata (Hegelm.) Hegelm. Locally abun- Aristida purpurascens Poir. dant in 2002-2003, edge of Sturgeon Pond (east) *Arthraxon hispidus (Thunb.) Makino (1988). CW sb, 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 351

CW f. Dominant on terrestrial portions of several Panicum clandestinum L. (1988). CW sb, CW f. islands. Panicum columbianum Schribn. CW f. Bromus ciliatus L. Panicum commutatum Schultes *Bromus inermis Leyss. CW f. Panicum dichotomiflorum Michx. (1988). RM sb, CW *Bromus japonicus Thunb. ex Murr. sb, CW f. *Bromus racemosus L. Panicum dichotomum L. *Bromus sterilis L. CW f. Panicum latifolium L. (1988). CW f. *Bromus tectorum L. CW f. Panicum stipitatum Nash. (1988, Schuyler). Calamagrostis canadensis (Michx.) Beauv. CW f. Panicum verrucosum Muhl. CW f. *Cenchrus tribuloides L. Panicum virgatum L. (1988). RM f, CW f. Cinna arundinacea L. (1988). RM sb, RM f. Paspalum laeve Michx. Critesion jubatum (L.) Nevski. (1988). CW f. Paspalum setaceum Michx. var. muhlenbergii (Nash) *Dactylis glomerata L. (1988). CW f. D.Banks Danthonia spicata (L.) Beauv. ex Roemer & Schultes. Phalaris arundinacea L. (1988). RM sb, CW sb, RM CW f. f, CW f. Deschampsia flexuosa (L.) Trin. *Phleum pratense L. [Abbott]. Digitaria filiformis (L.) Koel. CW sb, CW f. Phragmites australis (Cav.) Trin. ex Steud. (1988). *Digitaria ischaemum (Schreb. ex Schweig) Schreb. RM sb, CW sb, RM f, CW f. Present at CW in ex Muhl. CW sb, CW f. 1994; cover increasing. [Abbottϩ, ‘‘feathery *Digitaria sanguinalis (L.) Scop. (1988). CW sb, CW f. reed’’]. *Echinochloa crusgalli (L.) Beauv. (1988). RM sb, *Poa annua L. CW sb, CW f. [Abbottϩ]. *Poa compressa L. (1988). Echinochloa muricata (Beauv.) Fern. CW f. Poa palustris L. Echinochloa walteri (Pursh) Heller (1988). RM sb, *Poa pratensis L. CW f. CW f. *Poa trivialis L. (1988). RM sb, CW sb, RM f, CW *Eleusine indica (L.) Gaertn. (1988). CW f. f. Pseudo-viviparous plantlets noted on green- Elymus hystrix L. (1988). house seed-bank specimen. [Abbott]. Elymus villosus Muhl. ex Willd. Schizachyrium scoparium (Michx.) Nash (1988). CW f. Elymus virginicus L. (1988). *Setaria faberi Herrm. (1988). CW f. *Elytrigia repens (L.) Desv. ex Nevski. CW f. *Setaria pumila (Poir.) Schultes (1988). Eragrostis capillaris (L.) Nees Setaria viridis (L.) Beauv. CW f. *Eragrostis cilianensis (All.) Hubbard. CW f. Sorghastrum nutans (L.) Nash (1988). CW f. *Eragrostis curvula (Schrader) Nees. CW f. Spartina pectinata Link (1988, Schuyler, herbarium Eragrostis hypnoides (Lam.) BSP. (1988). CW f. only). Eragrostis minor Host Torreyochloa pallida (Torr.) Church (1988, Schuyler). Eragrostis pectinacea (Michx.) Nees. CW sb, CW f. Tridens flavus (L.) A.S.Hitchc. (1988). *Eragrostis pilosa (L.) Beauv. CW sb, CW f. *Vulpia myuros (L.) K.C.Gmel. var. myuros Eragrostis spectabilis (Pursh) Steud. (1988). CW sb, Zizania aquatica L. var. aquatica (1988). RM sb, RM CW f. f, CW f. [Abbott]. *Festuca elatior L. Festuca obtusa Biehler Pontederiaceae Festuca ovina L. Heteranthera multiflora (Griseb.) Horn. (1988). CW Festuca pratensis Huds. sb, RM f, CW f. Oberved at the Spring Lake spill- Festuca rubra L. (1988). CW f. way, at CW, and elsewhere. Appears to be more Glyceria canadensis (Michx.) Trin. CW f. frequent than H. reniformis. Glyceria melicaria (Michx.) F.T.Hubbard. CW f. Heteranthera reniformis Ruiz & Pavon (1988). RM f, Glyceria obtusa (Muhl.) Trin. CW f. Glyceria septentrionalis A.S.Hitchc. (1988). Pontederia cordata L. (1988). RM sb, CW sb, RM f, Glyceria striata (Lam.) A.S.Hitchc. (1988). RM sb, CW f. [Abbott] [Shoemaker]. RM f, CW f. *Holcus lanatus L. CW f. Potamogetonaceae Leersia oryzoides (L.) Swartz (1988). RM sb, CW sb, *Potamogeton crispus L. RM f, CW f. RM f, CW f. Potamogeton diversifolius Raf. (1988). [Schuyler]. Leersia virginica Willd. (1988). CW sb, CW f. Potamogeton epihydrus Raf. (1988). Leptoloma cognatum (Schultes) Chase. CW f. Potamogeton pectinatus L. (1988). *Lolium multiflorum Lam. CW f. Potamogeton pusillis L. CW f. *Lolium perenne L. (1988). CW f. Smilacaceae Melica mutica Walter Smilax glauca Walt. *Microstegium vimineum (Trin.) A.Camus. (1988). Smilax pseudochina L. Floodplain along Duck Creek. RM sb, CW sb, RM f, CW f. Common in some Smilax pulverulenta Michx. (1988). RM f, CW f. S. locations in CW, e.g., D. Cross Is. under woody pulverulenta var. pulverulenta. near Sturgeon canopy in 2004 where it is apparently replacing Pond, identified by D. Snyder. Cyperaceae. Smilax rotundifolia L. (1988). RM f, CW f. [Abbott] *Miscanthus sinensis Anderss. [Shoemaker]. Muhlenbergia schreberi J.F.Gmel. CW f. Panicum acuminatum Swartz. CW sb. Sparganiaceae Panicum anceps Michx. (1988). Sparganium americanum, Nutt. (1988). Panicum capillare L. CW sb, CW f. Sparganium eurycarpum Engelm. (1988). CW f. 352 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Typhaceae *Cleome spinosa L. Typha angustifolia L. (1988). RM f, CW f. [Shoemak- er]. Convolvulacee Typha latifolia L. (1988). RM sb, CW sb, RM f, CW *Ipomoea coccinea L. f. [Abbott] [Shoemaker]. *Ipomoea purpurea (L.) Roth Typha x glauca Godr. FWC. CW f. Identified by A.E. Cucurbitaceae Schuyler in 1996. *Citrullus colocynthis (L.) Shrad. Waif. Zannichelliaceae Polygonaceae Zannichellia palustris L. *Fagopyrum sagittatum Gilib. (1988, Blicharz). CULTIVATED EPHEMERALS: Araceae Solanaceae *Pistia strateoites L. Spring Lake; did not persist. *Petunia X hybrida Vilm. Asteraceae Poaceae *Tagetes erecta L. *Secale cereale L. *Zea mays L. Cannabanaceae *Cannabis sativa L. Planted and escaped (?), three lo- Pontederiaceae cations. *Eichhornia crassipes (Martius) Solms-Laub. Spring Lake several years; flowering observed but did not Capparidaceae overwinter.

Appendix 2. Plants noted by Bonaparte (1824), Abbott (1886, 1887, 1892, 1897, 1898), and by Shoemaker (1964) that are not presented in Appendix 1. The lack of an author citation means that there is no current synonym in Rhoads and Klein (1993), Gleason and Cronquist (1991), nor in Gray (1887). For the Abbott (1887) species, as in Appendix 1, no date is given; if they were listed in Leck et al. (1988), the date is indicated. Other Abbott citations are [Abbottϩ]. Each author mentions other species for which the specific name cannot be determined (e.g., Cyperus or Lemna); those are omitted. Taxonomy reflects Rhoads and Klein (1993) if possible. A (?) for certain Bonaparte species relates to difficulties interpreting hand written entries.

Bonaparte (1824): Andromeda racemosa (?) Perhaps this is Leucothoe racemosa (L.) A. gray, which is noted by Abbott (1898). Aquilegia canadensis L. Growing on rocks of the grotto at Point Breeze (2004), but not observed elsewhere. [Abbottϩ]. Aster patens Ait. Betula papyrifera Marshall Castanea vesca Cornus stricta Lam. Crateagus coccinea L. Erangianus americanus (?). Perhaps Fraxinus americanus. (‘‘A small one was located in front of the Lake House and a superb one at old Point Breeze’’ (Old Point Breeze was the first Bonaparte mansion that burned down; a second was built a short distance away). Euphorbia specananlia (?) Fagus sylvatica Hypoxis erecta L. In Gray (1887), possibly H. hirsuta (L.) Cov. Juglands porcina. In Gray (1887) Carya porcina Nutt. is pignut hickory, which in Gleason and Cronquist (1991) is Carya glabra (Miller) Sweet). Lycium carolinianum (?) Myosotis scorpioides L. [Abbottϩ]. Oxalis corniculata L.—‘‘Great quantity in grass, in woods, and around the aviary.’’ Philadelphus inodorus L. Potentilla canadensis L. Only this species is mentioned in Gray (1887), but Gleason and Cronquist (1991) would suggest this to be P. simplex Michx. ‘Great quantity in woods’. Pyrus coronaria L. Quercus aquatica Walter. In Gray (1887) this is Water Oak, which is now Q. nigra L. [Shoemaker]. Quercus tinctoria Gray. In Gray (1887) this is Scarlet Oak, which is now Q. coccinea Muenchh. [Shoemaker]. Robinia viscosa Vent. Rosa parciflora Perhaps Rubus parviflorus Nutt., thimbleberry. Rubus procumbens (?) Salix lareagailla (?) Salvia officinalis L. Cultivated for the kitchen. Saxifraga virginica. Probably S. virginiensis Michx. [Abbottϩ]. Spirea opulifolia. Cultivated? Syringa persica L. Cultivated. Syringa vulgaris L. Cultivated. Viola palmata L. Viola pedata L. 2005] LECK AND LECK: PLANTS OF A TIDAL FRESHWATER WETLAND 353

Appendix 2. Continued.

Abbott (authorities generally are those provided by Abbott (1887); others are from Gray (1887)): Amelanchier canadensis (L.) Medic. [Abbottϩ] Anemone nemorosa [Abbottϩ]. Aureolaria flava (L.) Farw. var. flava (Abundant enough to pick an armful). Brasenia schreberi J.F. Gmel. Old Mercer Co. record (Hough 1983). [Shoemaker]. Castanea pumila (L.) P. Mill. Celastrus scandens L. Clematis virginiana L. (1988) Cydonia vulgaris (Quince). [Abbottϩ]. Juglans cinerea L. Leucothoe racemosa (L.) A. Gray [Abbottϩ]. Muscari botryoides (L.) Miller. [Abbottϩ]. Polemonium caeruleum L. [Abbottϩ]. Populus diletata (ϭP. nigra L.). [Abbottϩ]. Prunus americana Marshall Prunus spinosa (‘‘Wild plum’’) [Abbottϩ]. Rhus venenata (ϭToxicodendron vernix (L.) Kuntze) [Abbottϩ]. Ribes grossularia (ϭR. uva-crispi L.) [Abbottϩ]. Rosa virginiana P. Mill. Rubus canadensis L. Rubus villosa Ait. In Gray (1887) called Common Blackberry, which is now R. allegheniensis T.C. Porter. Sabatia angularis (L.) Persh Salix purpurea L. Plantation noted. Sanguinaria canadensis L. [Abbotϩ]. Sisyrinchium burmudiana Gray (ϭS. anceps Cav.) [Abbottϩ]. Thalictrum anemonoides Michx. (ϭAnemonella thalictroides Spach. ϭ T. thalictroides (L.) Eames & B. Boivin) [Abbottϩ] [Shoemaker]. Trichochloa capillaris (‘‘Meadow comb grass’’). Uvularia perfoliata L. Viola cucullata Aiton. [Abbottϩ]. Viola spp. (White, yellow, purple) [Shoemaker]. Vitis labrusca ϫ riparia Shoemaker: Actaea pachypoda Ell. Hough (1983) notes this as rare on the inner coastal plain. Baptisia tinctoria (L.) Vent. Euphorbia marginata Pursh Iris prismatica Pursh Solidago odora Ait. Old Mercer Co. record (Hough 1983). Viola ricolor L. 354 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 132

Appendix 3. Species planted at the Duck Island constructed wetland between 1993 and 1995 by New Jersey Department of Transportation. Species designated with an * were not previously found in the study area. Author citations for species occurring in Appendix 1 are not given here.

Herbaceous Woody Acorus calamus Acer rubrum Iris pseudacorus Amelanchier canadensis Iris versicolor *Aronia arbutifolia (L.) Leersia oryzoides Ell. Nymphaea odorata Aronia melanocarpa Peltandra virginica Cephalanthus occiden- Pontederia cordata talis Potamogeton pectinatus *Chionanthus virginicus L. Sagittaria latifolia Clethra alnifolia Scirpus americanus Cornus amomum (Schoenoplectus pun- Cornus sericea gens) Fraxinus pennsylvanica Sparganium eurycarpum Hibiscus moscheutos Vallisneria americana *Ilex glabra (L.) A. Gray Zizania aquatica Ilex verticillata Lindernia benzoin Liquidambar styraciflua Magnolia virginiana Nyssa sylvatica Platanus occidentatus Quercus palustris Sambucus canadensis Vaccinium corymbosum Viburnum dentatum *Viburnum lentago L. *Viburnum opulus L.